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PROCEEDINGS

OF THE

ROYAL SOCIETY

QUEENSLAND

FOR 1949

VOL. LXI.

ISSUED 30th DECEMBER, 1950

PRICE : TWENTY-FIVE SHILLINGS

Printed for the Society by
WATSON, FERGUSON and COMPANY, Brisbane

The Royal Society of Queensland

Patron :

HIS EXCELLENCY LIEUT. -GENERAL SIR JOHN D. LAVARACK, C.B.,
C.M.G., D.S.O., C. de G., K.B.E.

OFFICERS, 1949

President :

DOROTHY HILL, D.Sc., Ph.D., A.N.C., F.G.S.

Vice-Presidents :

Professor H. C. WEBSTER, D.Sc., Ph.D., F.I.P., F.R.M.S.
Professor M. F. HICKEY, M.A., M.B., B.S.

Hon. Treasurer :

DOROTHEA F. SANDARS, M.Sc.

Hon. Secretary :

MARGARET I. R. SCOTT, M.Sc.

Hon. Librarian :
BETTY BAIRD, B.Sc.

Hon. Editors :

S. T. BLAKE,, M.Sc,
G. MACK, B.Sc.

Members of Council :

O. A. JONES, D.Sc., E. M. SHEPHERD, B.E., A. L. REIMANN, D.Sc., Ph.D.,
J. H. SIMMONDS, M.Sc., M.B.E., Professor L. J. H. TEAKLE.

Trustees :

F. BENNETT, B.Sc., Professor W. H. BRYAN, M.C., D.Sc., and
E. O. MARKS, M.D., B.A., B.E.

Hon. Auditor :

L. P. HERDSMAN.

Bankers :

COMMONWEALTH BANK OF AUSTRALIA.

CONTENTS

Vol. LXI.

No. 1 — Presidential Address : Energy and the Future of Mankind.

By H. C. Webster, D.Sc., Ph.D., F.Inst.P. (Issued separately,
30th December, 1950)

No. 2 — Contributions to the Geology of Brisbane, No. 1 — Local
Applications of the Standard Stratigraphical Nomen-
clature. By W. H. Bryan, M.C., D.Sc., and O. A. Jones, D.Sc.
(Issued separately, 30th December, 1950)

No. 3 — Marine Insects. By I. M. Mackerras, F.R.A.C.P. (Issued
separately, 30th December, 1950)

No. 4 — A New Ergot from Queensland. By R. F. N. Langdon,
M.Agr.Sc. (Issued separately, 30th December, 1950)

No. 5 — Revision of Bregmaceros with Descriptions of Larval Stages
from Australasia. By Ian S. R. Munro, M.Sc. (Issued
separately, 30th December, 1950) ...

No. 6 — Additions to the Flora of Arnhem Land. By C. T. White.
(Issued separately, 30th December, 1950) ... ...

•^No. 7 — Heavy Mineral Beach Sands of Southern Queensland.

Part II. — Physical and Mineralogical Composition,
Mineral Descriptions, and Origin of the Heavy Minerals.
By A. W. Beasley, Ph.D., D.I.C., F.G.S. (Issued separately,
30th December, 1950)

No. 8 — F. M. Bailey : His Life and Work. By C. T. White. (Issued
separately 30th December, 1950)

Report of Council

Abstract of Proceedings

Changes in Membership

Pages

1-1 1

13-18

19-29

31-35

37-54

55-58

59-104

105-114.

v.-vi.

vii.-xi.

xii.

50*o9^0

PROCEEDINGS

OF THE

ROYAL SOCIETY

QUEENSLAND

FOR 1949

VOL, LXL

ISSUED 30th DECEMBER, 1950

PRICE : TWENTY-FIVE SHILLINGS

Printed for the Society by
WATSON, FERGUSON and COMPANY, Brisbane

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Vol. LXI., No. 1.

Proceedings of the Royal Society
of Queensland

Presidential Address

ENERGY AND THE FUTURE OF
MANKIND

By H. C. Webster, D.Sc., Ph.D., F.Inst.P.

(Delivered before the Royal Society of Queensland, 28th March, 1949.)

Of all the abstract concepts of science, there is none which can
compare in importance with the concept of energy. There is little need
for me to explain to this Society the significance of the term energy,
but I should perhaps remind you of the well-known forms which energy
takes, viz., the forms of light, heat, sound, electricity, and the even
more familiar mechanical forms of kinetic energy (the energy of motion —
exemplified by a moving bullet), and potential energy (the energy of
position— exemplified by a wound clock spring). Then, too, energy can
assume a chemical form, such as the energy contained in fuel ; and,
finally, we have of recent years received startling evidence of the existence
of atomic energy.

The various things that happen on the earth, all actions, whether
of man, animals or of plants, all involve a conversion of energy- from
one form to another. The explosion of an atomic bomb, the eruption
of a volcano, a lightning flash, all represent well-understood types of
energy conversion. At the other end of the scale, the ticking of a
watch, the uttering of a word, vision, hearing, even the reception of a
sensation and the thinking of a thought, all imply energy conversions.

In all these conversions, there is no new energy created and no
energy destroyed. The energy which was in the universe at the beginning
is still in existence ana no new energy has appeared. New forms appear
and old forms disappear, but the gain always balances the loss. This
law is undoubtedly the most important law of science. It is really this
law which gives significance to the idea of energy ; without it the concept
would be meaningless.

This law refers to the total energy in the universe, not the energy
actually contained in or on the earth. The earth’s energy is not neces-
sarily constant in quantity ; in fact it is almost certainly varying all the
time. Energy is being added to the earth by the light and other radia-
tions received from the sun (and to a less extent from other celestial
objects), and energy is being lost by invisible radiations and in other
ways. The gain and the loss nearly balance out, but usually there is
not an exact balance.

The standard of living of the human race, even its survival depends
on the energy possessed by the earth. But the possession of energy
alone is not sufficient. The energy must also be in a form capable of
conversion to other forms, that is the energy must be available. Without

fUl ir

2 PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND

such available energy nothing can be made to happen, not even the
minor actions necessary for living. Unfortunately, energy can easily —
too easily, in fact — become converted into a form where it is no longer
available. Energy in the form of heat, i.e., the energy associated with
temperature, is never completely available. Only that part of the heat
of any material object which corresponds to a difference in temperature
between it and its surroundings can in practice be converted into other
forms of energy. The hot gases in a motor car cylinder have available
energy ; when they are cooled to the temperature of the cylinder they
have none.

Unfortunately, energy has a tendency to become converted into
heat. All energy conversions carried out on the earth, with or without
our direction, result in a certain proportion of the energy going into the
form of heat. There is, so to speak, a heat tax on all conversions.
While at first this heat energy may be still partly in available form, the
processes of conduction, etc., soon equalize the temperatures and render
the energy unavailable and therefore virtually useless. One might think
that there would be a possibility of obtaining energy on account of the
temperature difference between the surface of the earth and the cold
empty space within the shadow of the earth. Actually, this temperature
difference does allow the heat energy of the earth and air one important
energy conversion, the conversion into infra-red radiation. This con-
version is the ultimate fate of the heat energy. As a result of it there
is a continuous emission of radiation into empty space ; this is the main
way in which the earth loses energy. We can scarcely contemplate
making use of the temperature difference to obtain other forms of energy,
since, apart from the difficulties involved, to do so would lower the
average air temperature and human life can only be maintained over a
certain very limited range of temperature.

We have then on the earth two sorts of energy, useful energy, that
is, energy available for the operation of our machines, available for
making things happen, and useless energy, that is, unavailable energy.
When useful energy becomes converted into useless energy we can speak
of it as becoming degraded or consumed, as it is no longer available.

The most important of the machines is, of course, man himself.
I have already mentioned that the slightest action of the body, breathing,
the beating of the heart, even the transference of sensation, all represent
energy conversions. A high heat-tax is imposed on all these conversions,
so that the processes of life inevitably result in the consumption of
energy. To maintain life, therefore, an intake of energy is necessary.
This intake is in the form of chemical energy associated with the food
we eat.

The amount of energy intake depends on the sort of life a man is
leading, but a representative value for the average rate is about 150
watts ; this means 3.5 kilowatt-hours each day. (These units are the
most familiar of the energy units ; a kilowatt-hour is the unit ordinarily
used in selling electrical energy- — 150 watts is about a fifth of a
horse-power.)

As a result of the generation of heat within the human body, par-
ticularly within the trunk, the temperature of the interior of the body
is, under most climatic conditions, higher than that of its surroundings.
This interior temperature is subject to a system of automatic controls.
These regulate the way in which the body loses heat, and thus maintain

ENERGY AND THE FUTURE OF MANKIND

the interior temperature closely constant. For example, if the tempera-
ture of the surroundings increases somewhat, certain mechanisms increase
the ease with which heat passes from the interior of the skin ; if the
surroundings become colder, they decrease it. (Other mechanisms also
are involved, but details do not concern us.)

Man is assisted in this adjustment of his temperature by the fact
that the average air-temperature at sea-level is not very far removed
from the temperature at which the human body functions. This air-
temperature depends on the amount of radiation received from the sun.
As the average temperature of the earth increases, the rate at which
it loses energy to empty space also increases, and since loss and gain
must roughly balance, the greater the amount of energy the sun provides,
the higher the temperature of the earth.

The actual air-temperature at any place may vary quite considerably.
Over a certain range of conditions man’s regulating mechanism can cope
with the situation, but towards the limits of this range the adjustment
involves considerable strain and discomfort. At the lower end of the
temperature range the wearing of clothing and the use of houses assist
materially in this adjustment. They even permit life under conditions
in which otherwise it would be possible only by undertaking continuous
strenuous muscular exercise ; such exertion, of course, increases the heat
evolution within the body.

In his quest for comfort man has resorted to other measures, more
important from the point of view of my discussion to-night. He can
produce in a limited region such as the room of a house, a modified
climate, hotter or colder than the external climate, as may be required.
This modification of climate always demands the consumption of energy,

the conversion of energy from a useful form into a form of heat
which is useless. The actual steps in this degradation will differ in
different cases, but it always occurs. The amount of energy consumed
depends not only on the temperature differences maintained, but also
on such things as heat insulation, etc. Considerable technical develop-
ment has been devoted to reducing this wastage of energy. Even now,
however, a man may consume more energy keeping warm on a winter
evening than he consumes as food during the day.

If man’s needs were limited to food and warmth his energy require-
ments would be relatively easily met. But modern man demands far
more. He requires to cook his food to render it more palatable ; in so
doing he may expend almost as much energy as the food itself represents.
The growing of his food is no longer a matter which occupies merely
his own muscular effort. He requires all sorts of implements, many of
them requiring additional sources of energy, particularly fuel, for their
operation.

To obtain these and other implements man occupies himself in
manufacturing, making not only implements, but also houses and the
attributes of comfort, and making amusements and luxuries. All manu-
facturing involves the consumption of energy. Energy is consumed at
the mine where the ore is obtained, energy is consumed at the smelters
where the metal is extracted, energy is consumed on the railways when
the metal is taken to the factory, and energy is consumed at the factory
itself and in the subsequent journey to the user. Mostly such energy is
the result of the burning of fuel, coming from the chemical energy of

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND

wood and coal and oil. But energy must also be provided to the men
who work in these occupations, energy to provide them with the necessary
food and warmth and the other requirements of modern civilization.

There is a link between the cost in money of a manufactured article
and the amount of energy consumed in making it. They do not
correspond exactly, for often there are other factors involved, but it is
possible to estimate roughly the comparative cost of articles if we know
the energy involved in making them. Of course we have to make the
machinery used in making the articles, and we must include part of the
energy-cost of the machinery when we estimate the energy-cost of its
product.

Among the most important products of man’s labour are the fuels
themselves ; coal and coal products, petroleum and its products, forest
products, etc. The relative energy-profit on such operations is a matter
of the greatest concern to any community. To find the relative energy
profit, we add together the fuel energy expended in mining, transport,
refining, etc., and the food, etc., expressed in terms of energy, required
by the personnel employed. Then we subtract the result from the
energy provided by the fuel. This gives the profit, and we can express
it as a percentage in the usual way. If the total cost per ton is, say,
3,000 kw.-hrs., and if the energy we obtained from a ton is, say, 10,000
kw.-hrs., the relative energy profit is 230 per cent. It is not a coincidence
that the United States, which has plenty of easily-won fuel, is the richest
country in the world. On the other hand, England’s post-war financial
difficulties are related to the increasing difficulty of winning coal in a
country whose best coal-seams have already been exhausted.

The manufacturing activities of a modern industrialized country
may consume energy at the rate of more than 1,000 watts per head of
population, compared with the 150 watts per head required as food.
This does not, of course, represent the whole requirements of the popula-
tion ; it requires energy for artificial lighting, for transport, for radio and
other entertainment. As the use of motor cars becomes more widespread,
as the devices for providing entertainment become more elaborate, the
demands on energy increase. A petrol consumption of 100 gallons per
annum, a quite modest figure in pre-rationing days, represents an average
energy consumption at the rate of around 500 watts.

Average energy consumption rates for a typical person are somewhat
as f dIIows : —

Food ... ... ... ... ... ... 150 watts

Warmth ... 200

Manufacturing ... ... ... ... 2000

Transport ... ... ... ... ... 200 ,,

Miscellaneous ... ... # ... ... 450

Total ... ... ... 3000 ,,

Total Daily Consumption — 72 KW-hrs.

My estimates are based on peace-time requirements. In time of
war, energy is consumed at a rate many times greater than in peace,
and with tfie introduction of new weapons the consumption rises hugely.
The dropping of a single atomic bomb each day alone corresponds to a
consumption at the rate of forty thousand million watts, about 20 watts
per head of the world’s population.

If the spirit of man throughout the world is to be freed from the
chains of poverty, drudgery, and discomfort, if the standards of luxury

ENERGY AND THE FUTURE OF MANKIND

enjoyed among communities such as ours are to be shared by all mankind,
the average consumption of energy, food and fuel must inevitably
increase, and increase by a considerable factor. If war, disease and
famine decrease their toll, as we hope will be the case, the number of
energy consumers will also increase ; again we will require more energy.
How then are we to maintain and increase our present supplies of
available energy ?

Let us first examine the sources from which we obtain the energy
we consume at the present day. The most important group of these
sources, and the only absolutely indispensable one, is the food supplies.
We eat many things, some animal, some vegetable. Since the animals,
however, depend on vegetation as their source of food, we can regard
vegetation as the ultimate source of all our food. We must remember,
of course, that the energy we obtain from eating the flesh of animals is-
but a very small fraction of the energy those animals have consumed.

The growth of plants represents, in general, a storage of energy,
This energy is obtained from the light which the plant receives from the
sun by a photosynthetic process. This most important process occurs
chiefly, if not entirely, in the green leaves of the plant. To be precise,
it occurs in the chloroplasts , organs which contain the pigment chlorophyll
together with other pigments which may or may not participate in the
process. Under the influence of the light absorbed in the chloroplasts,
carbon dioxide is synthesized into sugar and energy is thereby stored
as chemical energy. This is the principal photosynthetic process, but
others also occur with which I shall not deal in detail. The overall
efficiency of the process is unfortunately very low. It has been estimated
that, of the solar energy falling on a green leaf, only two-thirds of one
per cent, is actually stored as chemical energy.

Production of Useful Energy.

Mechanism

Uses that part of the Sun’s
Energy which falls on

Efficiency
of use

Plants (photosynthesis in chloroplasts
containing chlorophyll) ...

Green leaves and green vegeta-
tion generally

0.66%

Hydro-electricity (rain on mountains)

Ocean and other water

0-001% (?)

Winds (heating of tropical regions) ...

All earth

perhaps

0.00001%

Photo- voltaic effect

Photo-voltaic cells ...

0.3%

The efficiency is not the same for all parts of the light spectrum,
though this seems to depend to some extent on the species of plant
involved. The energy of the infra-red radiation, which makes up about
half of the energy in sunlight, is not stored at all by plants.

Plants are of many kinds, not all of which contribute to our food
supplies either directly or indirectly. Among the non-food-producing
plants, however, are many which can still be regarded as useful ; for
constructional timber, for example, or for fuel. The fuels we are using
at the present day, chiefly wood, coal and oil, were all derived from
vegetation which, in the past, grew with the aid of sunlight. Coal and
oil have suffered many chemical changes but have still preserved the
chemical energy given to them by the sunlight. Oil may, in fact, have

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND

acquired additional energy, derived perhaps from the residuum of the
large heat energy which the earth possessed before it became a cold star.

In spite of the low efficiency of the photosynthetic process, it is of
vital importance, since it represents virtually the only one which is
replenishing those stocks of available energy which we are at present
using in such a prodigal fashion. Needless to say, the rate of exploitation
far outreaches the rate of restoration ; I doubt if there is any country
in the world which is actually increasing its reserves of energy in the
form of food and fuel.

These considerations bring home to us the seriousness of the tragedy
involved in the loss of arable land owing to wasteful farming transforming
it into a desert or dust-bowl. This has been stressed in relation to the
growing shortage of food — of which Sir John Boyd-Orr rightly warned
the peoples of the world — but this food shortage is but one aspect of
the more serious problem. Possibly, by using some of the areas at
present devoted to forests for food production, the food position could
be corrected, at least for some years, but this would aggravate the
general energy shortage. On the other hand, if ample supplies of dis-
posable energy become available, if the general energy problem is solved,
there would probably be no need for anxiety regarding the food position,
for I have little doubt that organic chemists will be able to discover
efficient processes (perhaps some new photosynthetic processes) whereby
foodstuffs can be made from carbon dioxide in factories in much the
same way as they are now made naturally in plants.

Incidentally, calculations which have been made of the extent of
coal and oil reserves in various countries may be misleading, for upon
the completion of exploitation of the richer and more accessible deposits
the energy costs of mining and transport will rise, perhaps sharply.
Without a serious drop in the standard of living, exploitation of the
remaining deposits may thus be impracticable. It is well known that
for technical and economic reasons, few coal-seams are ever completely
removed, and the cost of removing the residues at a later date may
well be prohibitive.

Fortunately, we do not depend entirely on fuel for the energy we
require for domestic heating, manufacturing and transport. A second
process initiated by the solar radiation provides us with a second source,
that of hydro-electric energy. The primary effect of the solar radiation
in this case is the evaporation of water, principally from the surface of
the oceans, but also from moist land, lakes, etc. The air thus moistened
may be carried by the winds, which are themselves a product of solar
radiation, over mountains and highlands and there, by cooling, the
moisture is deposited as rain. The water collecting at the high altitude
possesses energy, potential energy, and by suitably directing the water-
stream as it flows towards the sea, we can convert some of this potential
energy into other useful forms. In modern times, electricity is usually
the form of energy produced.

This energy is not produced without cost, i.e., without an initial
energy-outlay. Energy must be used in constructing dams, canals, pipes,
turbines, dynamos, etc. Most of the hydro-electric schemes which have
been installed in different parts of the world, however, have proved
highly profitable undertakings. The energy-cost of construction has been
covered by the energy produced within a relatively small number of
years. Probably there are still many possibilities for highly profitable

ENERGY AND THE FUTURE OF MANKIND

installations of hydro-electric schemes throughout the world, and many
others which, with care, would eventually return a profit in energy,
but only after many years.

Unfortunately there are conflicting demands on the available
streams of water in many countries, particularly in Australia. We
have frequently to choose between the utilization of the water for
stimulating an increase in food-production, thereby employing usefully
more of the sun’s radiation, or obtaining electric power from it directly.
Sometimes it is very difficult to determine which of these alternatives
will give the greatest overall energy-profit.

I have tried to obtain an estimate, for comparison with the plant-
growth method of utilizing solar radiation, of the average overall efficiency
of the production of electric power by evaporation from the oceans.
I can find no published figures, but on very rough assumptions, I arrive
at a figure of one part in 100,000 of the energy falling upon the ocean
being potentially convertible into hydro-electric energy. This is probably
a considerable over-estimate. Continental Australia with its low rainfall,
and small areas of high lands, is rather worse off than most countries
in relation to its size, as far as possibilities of hydro-electric generation
is concerned.

Other -means of utilizing solar energy have also been used to a
limited extent. Perhaps the most important of these is the use of
wind-power which was developed at one stage in the earth’s history to
a considerable degree, but recently tending to be abandoned on account
of its unreliability. It is extremely difficult, in fact impossible, to
estimate the overall efficiency of the wind-power method of using solar
energy, but it must be extremely low.

An interesting method for converting solar radiation into available
energy which has been suggested is the application of the photo-voltaic
effect. In the photo-voltaic cell, a comparatively simple electrical device,
electrical energy is generated when energy in the form of light falls on
the cell. Photo- voltaic cells are in fairly general use as illumination-
meters, photographic exposure-meters, and so on. It has been estimated
that the overall efficiency of the ordinary selenium photo-voltaic cell is
about one-third of one per cent. This method of utilizing energy would
thus be only about half as efficient as the utilization by plants. It
would have the advantage, however, that a supply of water, salts, etc.,
would not be needed, so it might be quite convenient for use in desert
and semi-desert areas, such as exist in parts of this country. It has
been estimated that 4,000 watts of power could be obtained from an
acre covered with such cells. Unfortunately, the cost in energy of the
manufacture of the cells is so high that it would be quite uneconomic
to proceed with large-scale projects on this basis.

Claims have been made by certain Russian investigators that a
much more efficient form of photo-voltaic cell has been discovered.
If these claims are substantiated, the invention may prove a very valuable
one, especially to countries like Australia. Maybe in the future, when
ruthless agriculture has denuded our mountains of all but solid rock
and converted our plains into deserts, the countryside will be covered
with photo-voltaic cells instead of forests, and maintenance engineers will
take the place of tillers of the soil.

It should not be thought, of course, that the methods already known
for utilizing the sun’s energy are necessarily the only or even the best

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND

methods available. The total average rate at which energy is received
from the sun works out at about two hundred million watts per head of
population. Compared with this figure our most extravagant require-
ments appear trivial. Our difficulties arise only from the extraordinary
inefficiency of all our methods of using sunlight. The situation is actually
somewhat worse than would appear from what I have said, for only a
small fraction of the sunlight actually falls upon green vegetation ; only
a part of the potential hydro-electric power can actually be obtained
because of cost of installation, and so on.

It is perhaps surprising that so few deliberate searches are being
carried out for new methods of converting solar radiation into com-
mercially useful energy. The reason is the absence of any obvious lead,
save the photo-voltaic cell scheme I have already mentioned. Other
possible schemes, such as the use of the thermo-electric effect, for example,
have been shown to be completely unprofitable. In the circumstances,
the solution of the problem is more likely to arise out of discoveries in a
completely unrelated field of physical or biophysical research than from
the results of investigations designed specially to. this end.

At the present time, the problem of the world’s energy deficiency
is being tackled along rather different lines. To understand this work,
we must enquire into the actual source of the solar radiation ; we must
determine why the sun retains its temperature, in spite of the enormous
amount of energy it is continually pouring out, mainly into the unbounded
vastness of interstellar space. The rate at which the sun loses energy
is nearly four hundred quadrillion watts (4 x 1026 watts), and some
process of energy-conversion must necessarily be occurring within the
sun for such an emission to continue without decreasing temperature.

It now seems fairly certain that the sun derives its high temperature
from the continuous conversion of its atomic energy into heat. The
existence of atomic energy has been recognized for a comparatively few
years. Its nature can be approximately explained in the following way :

Atoms are the building-blocks out of which matter is built, but an
atom itself is built up of smaller bits, rather in the way that the solar
system is built. Most of these sub-atomic particles are unimportant for
our present purpose, but there is in each atom one nucleus which is, so
to speak, the real body of the atom. This nucleus possesses most of the
mass of the atom, and with the nucleus is associated a certain amount
of energy. This energy is termed atomic energy. Atomic energy is then
really nuclear energy. It is conceivable that a nucleus might go out of
existence, in some sort of catastrophic process, in which case the atomic
energy would be converted into another form, probably into radiation.
Naturally, the mass would disappear with the disappearance of the
nucleus. On modern views, mass is really a measure of total energy
and, if a nucleus or anything else loses energy, it loses mass in proportion.

Such catastrophic disappearance of a nucleus has never been detected.
We do know of cases, however, in which part of the nuclear energy
becomes converted into other forms, and consequently the mass, the
energy indicator, becomes reduced. For example, it can happen when
a nucleus splits up into two separate nuclei ; the atomic energies
associated with two separate nuclei, added together, being in certain
cases less than the atomic energy associated with the single nucleus.
This disintegration process can occur spontaneously in radioactive

ENERGY AND THE FUTURE OF MANKIND

elements such as radium ; in fact, the value of radium as a method of
treating diseases is closely bound up with its ability to disintegrate and
thereby set free some of its atomic energy.

In the atomic bomb also there is a conversion of atomic energy due
to the splitting-up of nuclei ; in this case the nuclei of plutonium. This
reaction is not spontaneous, like the disintegration of radium, and
consequently we can control its initiation.

The process occuring in the sun is of quite a different nature.
Although the splitting-up of heavy nuclei, such as those of plutonium
and radium, can lead to the conversion of atomic energy into other
forms which can be used, the splitting-up of many light nuclei, notably
the breaking-up of a helium nucleus into four hydrogen nuclei, actually
involves the production of some atomic energy out of other forms of
energy. (Atomic scientists will realize, of course, that the manufacture
of hydrogen nuclei from a helium nucleus actually implies more than
a mere splitting-up, but I do not want to complicate the argument.)
If we reverse the process by building up helium from hydrogen, it should
be possible to set free some atomic energy, that is, convert it into other
forms which we can use. It may perhaps seem paradoxical that while
in one. case disintegration lowers atomic energy, in the other case it
leads to an increase. Nevertheless, by considering the structure of
nuclei in detail it is quite possible to arrive at a consistent explanation.
However, this is too long a story to enter into now.

This synthesis of helium from hydrogen is, we believe, continually
operating in the sun. It is scarcely feasible that this synthesis should
occur simply through four hydrogen atoms coming together. Rather it
would appear to take place in stages, a carbon nucleus acting as an
intermediary in the process. The details are. still somewhat speculative.
The present theory postulates a chain of six nuclear reactions, which
incorporate the hydrogen nuclei one at a time and set the carbon nucleus
free again at the end, the net result being the combination of the four
hydrogen nuclei into helium. We can equally well regard the process as
beginning with nitrogen instead of carbon, but this is a matter of detail.

Adopting this theory of the sun’s activity, it is not difficult to provide
a reasonable account for the approximate constancy of the sun’s tempera-
ture and energy output. -I say approximate because disturbances of the
sun’s surface, notably sunspots, eruptions, etc., are of comparatively
frequent occurrence, and appear to influence the amount of radiation.
It seems likely that the sun’s output will change relatively slowly over
the next few millions of years. It will probably slowly increase at first
but, after the lapse of many millions of years, it will drop and continue
to drop until the sun is a cold star like the earth and all life is extinct.
It is possible, of course, that at some earlier stage the sun will explode,
as some stars have been known to do, and life would then be destroyed
in a more sudden and spectacular manner. The sun is, after all, a
large-size atomic bomb.

The energy that results from the synthesis of helium is very great,
far greater than any ordinary burning of a comparable mass of fuel
can produce.

The energy-output of the sun is obtained at a cost of less than a
quarter of an ounce of hydrogen per thousand kilowatt-hours. The
total consumption is large, amounting to three thousand billion tons
(3 X 1015 tons) each year, but this is only about one billionth part of the

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND

Energy Derived from Fuels.

Fuel

Form of Energy-
Utilized

Energy Obtained
(KW-hrs per Kgm.)

Wood

Chemical

say 5

Coal

Chemical

say 10

Oil

Chemical

say 12

Plutonium

Atomic

say 30,000,000

Uranium-235

Atomic

say 25,000,000

HYDROGEN

Atomic

150,000,000

sun’s total mass. We do not know precisely what proportion of the
sun consists of hydrogen, but there is evidence that hydrogen must
constitute an appreciable fraction of the total, so there is no need to
fear the hydrogen supply running low for a few million years or so.

Since the earth, like the sun, is composed partly of hydrogen —
about one per cent, of the earth’s crust consists of this element — one is
led to consider whether there is any chance of setting-up, on the earth
and under our control, a machine in which hydrogen, in relatively small
quantities, could be converted into helium. Such a process could not
be made to occur spontaneously, for there are decisive factors which
prevent this, but it is conceivable that some process might be devised.
We should want one different from the solar process ; it could not be
considered because of the enormous temperatures involved. We want
then some other process leading to the synthesis of helium from hydrogen.

Such a process, if successful, might provide us with all the energy
we need at a comparatively small cost in hydrogen. Taking a figure of
ten thousand million kilowatts as the outside estimate of our demands,
this corresponds to a consumption of somewhere about a ton of hydrogen
per day. As the oceans alone contain about a hundred thousand billion
tons (1017 tons) of hydrogen, this consumption could scarcely be regarded
as excessive. In fact, nearly as much hydrogen is probably being wasted
at the present time by the escape of hydrogen gas resulting from the
electrolysis of water.

Naturally we could hardly expect 100 per cent, efficiency from our
machine, i.e., we could hardly expect all the atomic energy-reduction to
appear as useful energy. Further we should have to employ a consider-
able amount of useful energy in making the machine itself. Even if
we had an overall efficiency of only one per cent., however, the con-
sumption of hydrogen could scarcely be regarded as serious.

You may wonder why, when we have a source of energy ready to
hand in the plutonium bomb and the nuclear fission pile used in making
it, I have stressed the importance of the hydrogen process. The fact
is that while the first development of atomic energy machines, using
uranium and thorium as the raw materials, may provide an immediate
solution to the pressing problems of fuel shortages, it cannot be regarded
as anything but a temporary solution. This is due to the fact that the
high-grade ores of uranium and thorium will almost certainly be rapidly
exhausted ; in fact, the atomic bomb manufacturing programme of the
United States will probably exhaust them before the end of the century.
Low-grade ores may still be used — they almost certainly will be used
for atomic bombs unless a better bomb is invented in the meantime —
but their use will not be profitable in terms of energy. Precisely at

ENERGY AND THE FUTURE OF MANKIND

what stage the process will cease to provide an energy-profit cannot
of course be predicted.

If the long-term solution must lie in a hydrogen-helium process,
how are we to discover a suitable process ? There is no obvious line of
attack. For this reason the Atomic Energy Establishments of Britain,
France and the United States, together with University and other
research laboratories, are devoting their activities very largely to quite
general researches into nuclear physics. Only by the process of slow
compilation of information concerning nuclei and their behaviour, only
by the elucidation of their fundamental properties and the phenomena
connected with them, can we hope to make progress. The solution,
when it comes, is more likely to result from some apparently quite
irrelevant research than from a straight-forward attack on the problem.

This is the reason why physicists are impressed with the importance
of nuclear research ; this is the reason why they are sometimes somewhat
impatient of the apathy, even obstruction, with which their proposals
are often received. Those who, because of the belief that nuclear research
necessarily means military research, or for personal or political advantage,
oppose or obstruct nuclear research, are doing a very real disservice to
mankind. If all the peoples of the world are to possess and maintain
a standard of comfort and luxury such as the more fortunate peoples
now enjoy, the energy supply problem must be solved, and must be
solved soon.

Vol. LXI., No. 2.

CONTRIBUTIONS TO THE GEOLOGY
OF BRISBANE

No. 1. — Local Applications of the Standard Stratigraphical

Nomenclature.*

By W. H. Bryan, M.C., D.Sc., and O. A. Jones, D.Sc.
University of Queensland.

(Received 17 th May, 1949 ; read before the Royal Society of Queensland ,
31s£ October, 1949 ; issued separately 30 th December, 1950).

In the following proposals an attempt has been made to conform
to the Australian Stratigraphical Nomenclature suggested recently by
Glaessner and others (1948). In accordance with rule III f of the Code
as there set out the new names now introduced are explicitly defined,
the geographical features from which the names were taken are stated
and the specific type localities cited. Where changes are proposed the
reasons are concisely stated.

Rocksberg Greenstones. — A formational name introduced to
replace the term “ Greenstone Series ” of Denmead (1928). The forma-
tion consists almost entirely of metamorphosed andesitic and basaltic
lavas and tuffs. The name is taken from the village of Rocksberg, near
Caboolture, where Mr. R. T. Mathews, who is working on the formation,
reports it is typically developed.

The reason for the proposed change in name is the absence of certain
knowledge of its age and range in time, which precludes it from any
more precise category than that of a formation. It is now ranked as
such and named accordingly.

Bunya Phyllites. — A formational name introduced to replace the
term “ Bunya Series ” of Denmead (1928). The formation consists
essentially of pelitic rocks such as mica phyllites with some quartz-mica
schists ; psammitic rock types are well represented only in the eastern
part of the area, and even there they are restricted to the uppermost
part of the formation. As here redefined, the formation excludes certain
cherts and quartzites (which were included in the uppermost part of
Denmead’s Bunya Series) and places the top of the formation immediately
below the lowest of these, the Kenmore Quartzite, which outcrops near
the mine at Gold Creek and can be followed in a direction S. 30° E. to
Fig Tree Pocket and thence in a more easterly direction to the Carrington
Rocks at Corinda. The formation conformably succeeds the Rocksberg
Greenstones, and is conformably overlain by the Neranleigh- Fern vale
Group. The name is based on Bunyaville, an outer suburb within the
area of Greater Brisbane, where the formation is well developed.

The reason for the proposed change in name is the absence of certain
knowledge of its age and range in time.

*For several years the authors have been collecting material for use in a book
to be published under the title o “The Geology of Brisbane.” Following a study
of the relevant literature and alter considerable work in the field they have come
to a number of conclusions that differ importantly from those now generally accepted.
These conclusions will be set out as such in the book, but it would seem that the
arguments on which they are based would be more appropriately stated in the
Proceedings of this Society.

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND

Neranleigh-Fernvale Group.— A new composite name which
incorporates in the one term both the “ Neranleigh Series ” and the
“ Fernvale Series ” of Denmead (1928) and at the same time changes
the category from series to group. As here redefined the Group includes
certain cherts and quartzites which were originally included in Denmead’s
Bunya Series and the lowest of which, the Kenmore Quartzite, is regarded
as the base of the group. The name Neranleigh was originally taken in
part from the village of Neranwood and in part from the town of
Beenleigh, both to the south of Brisbane, while the name Fernvale was
derived from a township in the Brisbane Valley to the west of Brisbane.
The type locality for the Group as such, now selected by the authors, is
the valley of Moggill Creek, within the area of Greater Brisbane.

The reasons for the amalgamation of these two “ Series ” are : The
absence of any structural break within the group and the recurrence of
similar lithological types throughout the group. For many years grey-
wackes had been regarded as characteristic of the lower (Neranleigh)
part of the group and radiolarian jaspers as equally typical of the upper
(Fernvale) part, but the Moggill Creek section shows some of the jaspers
occurring at relatively low horizons within the group and some of the
greywackes at relatively high levels. The group is highly variable
lithologically and includes, in addition to the greywackes and jaspers
(which have been over-emphasised in the past), such rocks as phyllites,
quartzites, both thin-bedded and massive, impure limestones and
calc-epidote rocks.

The Group conformably succeeds the Bunya Phyllites.

Neither the age nor the range of the Group is known sufficiently
accurately to enable the use of a more precise term than “ group.”

Brisbane Metamorphics. — A name introduced to replace the term
“ Brisbane Schists.” Although of uncertain origin the latter designation
has been widely used for many years as a comprehensive name covering
the immense thickness of metamorphosed marine sediments, tuffs and
lavas made up of the Rocksberg Greenstones, the Bunya Phyllites and
the Neranleigh-Fernvale Group as defined in the preceding paragraphs.
(See Bryan and Jones 1944, p. 13.)

The geographical portion of the proposed name is taken from the
city of Brisbane within and near which the Metamorphics are typically
developed.

The reason for changing the second part of the name from “ Schists ”
to “ Metamorphics ” is that although schistose rocks of several types
are present they are by no means as dominant as the original name
would suggest.

The term “ Brisbane Metamorphics ” is not in strict accordance
with the Stratigraphical Code, but the authors feel that some additional
and more comprehensive designation is warranted to indicate the unity
in general characters which distinguishes the Brisbane Metamorphics
from all later stratigraphical units and which overrides those less funda-
mental differences which have led to the recognition within the
Metamorphics of two distinct formations and one group. The term
“ Brisbane Complex ” was considered as an alternative, but was rejected
as being at odds with this essential unity and, moreover, as likely to
lead to confusion.

CONTRIBUTIONS TO THE GEOLOGY OF BRISBANE

Brookfield Volcanics. — A name proposed for a succession of
flows, tuffs and agglomerates of varied character but predominantly
rhyolitic. The name is taken from the village of Upper Brookfield in
the western part of Greater Brisbane. The Volcanics are typically
developed near this locality on top of the divide between Moggill and
Pullen Vale Creeks.

The age of the Brookfield Volcanics has not been determined but
they rest unconformably upon steeply dipping beds of the Neranleigh-
Fernvale Group.

Brisbane Tuffs. — This name represents a reversion from the term
“ Brisbane Tuff ” now in common use to Dunstan’s (1916) original
designation for the accumulation of tuffaceous materials of a rhyolitic
nature occurring within, but almost at the base of the Ipswich Coal
Measures as developed at many points within the city of Brisbane. The
Tuffs have been assigned to the Middle Triassic. ( See Bryan and Jones
1946, p. 52.)

The use of the plural is advocated as an indication that the tuffaceous
material is of several different kinds, including water-laid tuffs, wind-
blown tuffs and welded tuffs (Ignimbrites) , and that these do not all
occur on precisely the same stratigraphical horizon.

Ipswich Coal Measures. — It is recommended that this name be
selected from the several synonyms now in common use ( see Bryan and
Jones, 1944, p. 41) for the freshwater shales and sandstones, some of
them coal-bearing, that with conglomerates and some tuffs make up a
succession of 4,000 feet of strata, the lower limits of which occur on
the right bank of the Brisbane River near Mt. Crosby where they rest
unconformably on beds of the Neranleigh-Fernvale Group, and the
upper limit of which is immediately beneath the Aberdare Conglomerate
at Denmark Hill, Ipswich. They have been assigned to the Middle
Triassic ( see Jones and de Jersey 1947d, p. 82 ; Bryan and Jones 1946,
p. 54). The place-name is based on Ipswich, and the Measures are
typically developed within and to the north, east and south of that city.
With rather more precise knowledge of the range of these Measures, it
may be possible to promote them to a Series in the sense of the Code.

Bundamba Sandstones. — A formational name proposed in place of
the Bundamba Series of Cameron (1907), for coarse fresh-water grits and
sandstones, often showing cross-bedding with thin interbedded shales
which are commoner towards the base, near which one thin coal-seam
occurs. The base is marked by the Aberdare Conglomerate which
succeeded the Ipswich Coal Measures after a short erosion interval.
The Sandstones have been assigned to the Upper Triassic (see Bryan
and Jones 1946, p. 54). The place-name is based on an outer suburb
of Ipswich, where the Sandstones are typically developed.

The reason for the proposed change is that, in the absence of certain
knowledge of their age, these sandstones do not form a “ Series ” in
the sense of the Code.

Brighton Beds. — A name first proposed by Woods (1947), and
supported here, for fresh-water micaceous sandy shales often white in
colour, but sometimes biscuit brown, together with red and white sand-
stones and including a curious and easily recognizable horizon of oolitic
character. The beds are horizontal and the base and thickness are as

16 PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND

yet unknown. They have been assigned to Lower Jurassic ( see Jones
and de Jersey 1947b, p. 11), and are unconformably related to the nearby
Ipswich Coal Measures.

The place-name is taken from Brighton, near Sandgate, where the
Beds are typically developed.

Redbank Plains Formation. — A name introduced to replace the
term “ Redbank Plains Series ” of Jones (1927). The formation consists
of fresh-water clays, mudstones, shales and soft micaceous sandstones
together with interbedded basalts on 'several horizons. The formation

THE GEOLOGY OF BRISBANE

Comparison of Proposed Stratigraphical Terms with those now

COMMONLY IN USE.

European

Record

Names now in use

Names now proposed;

Recent

Lone Pine Gravel

Pinkenba Beds
Lone Pine Gravel

Pleistocene . . .

Pliocene

Miocene

Oligocene

Petrie Series

Petrie Formation

Eocene

Redbank Plains Series

Redbank Plains Formation

Cretaceous

Jurassic

Brighton Beds

Triassic

Bundamba Series
Ipswich Series
Brisbane Tuff

Bundamba Sandstones
Ipswich Coal Measures
Brisbane Tuffs

Permian

Carboniferous

Devonian

Silurian

Fernvale Series

Neranleigh-

Fernvale ^ o

Group § S

rD lT'

Ordovician . . .

§ 42 Neranleigh Series

aj c/3

r ^ . (H

•£ ^ Bunya Series

.<2 o

Bunya Phyllites ^ g

M-l

Cambrian

Greenstone Series

-4->

Rocksberg g

Greenstones

CONTRIBUTIONS TO THE GEOLOCxY OF BRISBANE

overlies the Ipswich Coal Measures unconformably and has been assigned
to the Eocene (see Bryan and Jones 1946, p. 67). The name is taken
from and the formation typically developed on the Redbank Plains,
near the township of Goodna.

The reason for the proposed change of name is that this succession
does not constitute a ” Series ” in the sense of the Code, the range in
time being as yet uncertain.

Petrie Formation. — A name proposed to replace the term Petrie
Series of Jones (1927). The formation consists of fresh- water ferruginous
quartzite-breccias, fine-grained micaceous white and red sandstones and
some oil-bearing shales. The formation rests with a slight unconformity
upon the Ipswich Coal Measures and has been assigned to the Oligocene.
(See Bryan and Jones 1946, p. 67.) The name is taken from the township
of Petrie, to the north of Brisbane, where the formation is typically
developed.

The reason for the proposed change in name is that the succession
does not constitute a “ Series ” in the sense of the Code, the range in
time being as yet undetermined.

The Lone Pine Gravel. — A name first proposed by Bryan (1938)
and supported here for semi-consolidated quartzitic conglomerates of
fluviatile origin found at relatively high levels on the margins of the
lower part of the Brisbane River. The gravel is of late Kainozoic age.

The name is based on a tourist resort on the left bank of the Brisbane
River some fifteen miles by water above the city, where the gravel is
typically developed.

Pinkenba Beds. — A name now proposed for semi-consolidated
sands, silts and sandy clays of marine and estuarine origin which are
well developed under, the low-lying flat areas about the mouth of the
Brisbane River. The Beds are of late Kainozoic Age.

The name is based on an outer suburb of the City of Brisbane, where
the Beds are typically developed.

LITERATURE CITED.

Bryan, W. H., 1938. — “ The Pebbles on my Garden Path.” Oueensl. Nat. 10,
83-93.

Bryan, W. H., and Jones, O.A., 1944. — “A Revised Glossary of Queensland
Stratigraphy.” Univ. Queensl. Papers, Dept. Geol. 2 (N.S.) No. 11.

— ,1946. — ‘‘The Geological History of Queensland.” Univ. Queensl.
Papers, Dept. Geol. 2 (N.S.) No. 12.

Cameron, W. E., 1907. — Second Report on the West Moreton (Ipswich) Coal-
field.” Geol. Surv. Pub. No. 204, 37 pp., 2 maps, 1 plate, 8 figures.

Denmead, A. K., 1928. — ‘‘A Study of the Brisbane Schists.” Proc. Roy. Soc.
Queensl., 39, 71-106, pis. vi-x and text-figs.

Dunstan, B., 1916. — “ Queensland Geological Formations. Appendix.” School
Geography of Queensland. G. Harrap.

Glaessner, M. F., Raggatt, H. G., Teichert, C., and Thomas, D. E., 1948. —
‘‘ Stratigraphical Nomenclature in Australia.” Aust. Jour. Sci. 11, (1) ,
pp. 7-9.

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND

Jones, O. A., 1927. — “ The Teitiary Rocks of the Moreton District, South-East
Queensland.” Proc. Roy. Soc. Queensl., 38, pp. 23-46, pis. 6-8, 4 text
figs. 2 maps.

Jones, O. A., and de Jersey, N., 1947a. — “ The Flora of the Ipswich Coal
Measures.” Univ. of Queensl. Papers, Dept.Geol., 3 (N.S.) No 3.

, 1947b. — “ Fertile Equisetales and other Plants from the Brighton

Beds.” Univ. Queensl. Papers, Dept. Geol., 3 (N.S.) No. 4.

Woods, J. T., 1947. — “ Stratigraphical Notes on the Brighton Beds.” Univ.
Queensl. Papers, Dept. Geol., 3 (N.S.) No. 4, pp. 12-16.

Vol. LXI., No. 3.

MARINE INSECTS*

By I. M. Mackerras, F.R.A.C.P., Queensland Institute of Medical

Research, Brisbane.

(Received 4th April , 1949 ; read before the Royal Society of Queensland,
31 st October, 1949 ; issued separately 30 th December, 1950).

Insects are highly successful and widely distributed animals, which
have established themselves in many environments and come to dominate
many ecological associations, and yet there are comparatively few records
of their occurrence in the seas. This review has been stimulated by
Wassell’s (1948) most interesting discovery of Pontomyia natans Edw.
in Australian waters and by a few observations we made on a coral cay.
It has been necessary to draw largely on information from other parts
of the world, because Australian records are for the most part scattered
in the literature and rarely accompanied by details of habitat or
behaviour.

THE EVOLUTION OF INSECTS.

It is necessary, in the first instance, to remember that insects
evolved as terrestrial animals (Tillyard, 1930 ; Tiegs, 1949). They arose,
apparently, in the Palaeozoic from primitive terrestrial Myriapods.
The Aptera came first, and Collembola have been found in the Devonian,
where they lived “ in peat bogs along with Acarids, Crustacea and the
most primitive types of vascular plants/’ a terrestrial, if damp, situation.
There was a great evolution during the Carboniferous ; winged insects
appeared, and the ancestors of many existing Orders became
differentiated. Progressive reduction in loss of water through the cuticle
was undoubtedly an important factor in this adaptive radiation, but
some insects, such as the stoneflies and the dragonflies, were already
becoming adapted to an aquatic existence during part of their life-cycle.

From this time, and still more in the Permian, representatives of
more and more Orders invaded the fresh waters of the earth, so that the
aquatic insect fauna became an abundant and varied one, which showed
many remarkable adaptations to life in ponds, streams and lakes. It is
from these that most, though not quite all, of the marine insects arose.

The insects that live in fresh waters are many, but they only
represent a portion of the Orders and a fraction of the families of all
the insects. So, too, the marine insects only represent a fraction — -
indeed, a small fraction — of the groups which have invaded fresh water.

ADAPTATIONS TO AQUATIC LIFE.

There are two quite different basic adaptations to aquatic life.
The first is shown by those insects which are insulated against the water,
frequently by means of a close pile of hairs or scales, the result being
that they are not actuclly in contact with the water. Water-beetles
and such bugs as the water-skaters are examples of this type. They
may run upon the water, they may even dive teneath the surface,
covered with a film of air and carrying their bubble of air for respiration,
but they are not wetted. They do, however, live in a saturated
atmosphere, and the film of air may play an important part in respiration
beneath the surface.

Presidential address to the Entomological Society of Queensland, 14th March, 1949.

20 PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND

The second group is truly aquatic. Their bodies are wetted, and
they cannot survive out of the water in the aquatic stage of their life-
history. They show many special adaptations, of which the most
important are : —

1. They no longer need protection from loss of water through
the body’s surface ; but they do need means to regulate ionic
exchange between their body-fluids and the external
environment.

2. They require special means to cope with respiratory exchange,
either by breathing-tubes which pierce the surface (or some-
times the tissues of aquatic plants), or by blood or tracheal
gills, or by increased cuticular permeability to dissolved gases.

3. As water exchange between the insect and its environment is
unrestricted, there is no need for any mechanism for storing
insoluble excretory products.

4. Wings are obviously useless impediments during the aquatic
stages of an insect’s life.

Marine insects show the same basic adaptations but they meet
special difficulties.

1. The atmosphere surrounding the hydrophobe insects is not
quite saturated, and therefore these insects may need more
efficient means to control evaporation from the body-surface
than their fresh-water relatives.

2. Whether on the surface or beneath it, marine insects face
conditions of turbulence which are not usually found in the
fresh waters except on the margins of large lakes. Even in
swiftly flowing streams, the inhabitants only have to align
themselves with the direction of flow, and do not need to cope
with changes in direction due to tides, currents, winds, and so
on. Buxton (1926) drew attention to this factor, and regarded
it as an important though not a vital one in limiting invasion
of the sea.

3. In addition to movement, bottom-dwelling insects must be
able to withstand changes in hydrostatic pressure due to the
rise and fall of the tide.

4. The turbulence-factor determines that insects living below the
surface cannot depend on respiratory tubes for their gaseous
exchange. Thus, it is well known that mosquito larvae cannot
survive in waters where there is splash and wave motion, and
it is significant that the larvae of marine Chironomidae have
cutaneous respiration and their pupae lack breathing trumpets.

5. Of even greater importance is the need to regulate the exchange
of water and ions, and to cope with a reversed osmotic gradient.
Morphological evidence of this factor is seen in the reduction
of anal papillae in mosquitoes and Chironomidae which live in
saline waters.

6. The food factor may also be important. There are so few
higher plants in the sea that invasion by phytophagous insects
would be extremely difficult. Those which live on Algae,
diatoms, etc., in fresh water find less violent change required,
and so also do those which live on animal food.

MARINE INSECTS

7. A further hazard is probably found in the predatory life. The
numbers and variety of predators one sees in ponds and streams
are impressive, but those one encounters on rocky foreshores
and coral reefs are even more impressive, and they differ, too,
in their methods of finding and seizing their prey.

Collectively these are formidable barriers. That they are real is
indicated by the fact that almost the only insects to become established
in the sea are hydrophobes, which skate on the surface, or shelter in
rocks or weeds when submerged, and the larvae of certain Diptera, which
have cuticular respiration and efficient means of hiding from their enemies.
It may be noted, too, that marine insects are nearly always small
(Miall, 1903).

Invasion of the sea probably occurred by two routes : gradually
from streams through their estuaries, and by more abrupt changes from
pools and swamps to the littoral zone and the reefs. The open ocean
appears to have been reached only once by each path. These lines are
indicated in Table 1, as well as the rapid decrease in the variety of
Orders as one proceeds seaward.

TABLE ].

Orders of Insects in which Aquatic Species are Known.

Order

Fresh

Estuarine

Littoral

Pelagic

Collembola (H)

Ephemeroptera (A)

Odonata (A)

(x)

Perlaria (A)

Hemiptera (H, A)

Coleoptera (H, A)

Hymenoptera (A)

x(b

Neuroptera (A)

Diptera (A)

Trichoptera (A)

Lepidoptera (A)

x(2)

Orthoptera (? H)

,x(2)

H = Hydrophobe adults (sometimes all stages when there is no metamorphosis).

A = Aquatic early stages.

(1) Parasitic on aquatic insects.

(2) A few genera only. Siphonaptera also occur on marine mammals.

ESTUARINE FAUNA.

The change from fresh water to salt in river estuaries is fairly
gradual, so one would expect to find an equally gradual decrease in the
fresh water insect fauna as one approaches the sea, and an associated
appearance of forms showing progressive adaptation to life in salt water.
Such an area should be one of evolutionary change and speciation.
Actually, this may not be true, and Buxton quotes evidence that in
saline lakes of Europe there is a critical salt concentration which bars
colonisation. In waters with a saline content of less than 2.5% species
of Odonata, Hemiptera, Neuroptera, Diptera and Trichoptera were
found, but only Diptera in those with a salt content greater than 2.5%.

It is difficult to compare these findings with what happens in estuaries
because few accounts of estuarine insect faunas have been available to me.
Lindberg (1937) gives detailed records of the Hemiptera and Coleoptera
of a Finnish bay opening into the Baltic. The salt content of the water

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND

was very low (from 0.1% to 2% at different stations), and both Orders
were well represented, the Hemiptera by the families Hebridae (1 species),
Gerridae (5), Mesoveliidae (1), Veliidae (1), Corixidae (7), Notonectidae (2)
and Nepidae (2), and the Coleoptera by Haliplidae (6), Dytiscidae (31),
Gyrinidae (5), Hydrophilidae (17), Dryopidae (1) and Chrysomelidae (1).
Only 10 species of beetles and 2 of bugs were taken in the most saline

(seaward) station. The Plymouth workers ( , 1931) record a

caddis fly (Leptocerus) at the top of the tidal region in the Tamar R.
We have seen Gerrids in the tidal part of the Mary R., and wondered
whether they indicated the path taken by the ancestors of Halobates
on their way to the sea. Chironomid larvae, apparently of two different
subfamilies, have been dredged from muddy bottoms at considerable
depth (e.g., Orthocladius oceanicus Pack, from 30 fathoms in Salem
Harbour, U.S.A.).

All this is very meagre. A large and interesting field clearly awaits
the naturalist, who will work in the lower reaches of our Queensland
rivers and make salinity records in parallel with his insect collections.

LITTORAL FAUNA.

The insect fauna of the shore-line is more varied and better known.
The types of situation also vary greatly, and we may consider five very
different environments separately.

Mangrove swamp.

It is difficult to know whether to class this environment as littoral
or estuarine. I have chosen the former because the Diptera at least
have extended to it from the pools and swamps of the land rather than
from the streams which flow past its edges. This is well shown by the
mosquitoes, nearly all of which can breed — and sometimes do so freely —
in water of all gradations from perfectly fresh to brackish or even con-
centrated sea water, for example Anopheles punctulatus farauti Lav. in
water from 0 to 1.4% and A. amictus hilli Edw. from 0 to 4.2% saline
content (Lee and Woodhill, 1944).

The best known insects of the mangrove swamp are Nematocerous
Diptera. Mostly they breed in the temporary pools of the inner mangrove
zone, left by the high tide and more or less diluted by rain or seepage.
Among the Australian mosquitoes, there are, in addition to the
Anophelines mentioned, Aedes vigilax Sk., A. alternans Westw.,
A. longirostris Leic., Culex ' sitiens Wied., C. vishnui Theo. and
C. fraudatrix Theo. (Lee, 1944). It is interesting to observe how com-
pletely these forms are restricted to situations which are cut off from
direct contact with regular tidal waters. Other occupants of the same
environment are Chironomidae (Chironominae) and Ceratopogonidae.
Adult Culicoides are extremely abundant, and have been found breeding
in mangrove swamps in other parts of the world, but the only larvae so
far found in this country live in fresh water (Marks, 1947).* The
Dolichopodid, Thinophilus wasselli Hardy, is plentiful on uncovered
tidal mud (Hardy, 1935).

* Since this was written, Lee (Aust. J. Sci ., 12 ; 74, 1949) has found the early
stages of a pest species in the Salicoinia zone above ths mangroves in New South
Wales.

MARINE INSECTS

Open salt marsh.

The insect fauna of the salt marshes in low-lying country behind
the sea-front illustrates the effects of salinity referred to by Buxton,
most of the species being restricted to brackish waters. It is, however,
richer than the estuarine fauna, doubtless due to freedom from tidal
action and better shelter from predators. In Hawaii, Williams (1936,
and later papers of the series) recorded water-beetles (Enochrus) , a small,
active Corixid bug, and occasionally a dragon-fly (Anax) from such
brackish waters. Lispine flies, Acalyptrates (chiefly Ephydridae) and
Dolichopodidae frequent the margins of the pools.

In Australia, several of the mosquitoes mentioned above breed also
in salt marshes. In addition, Aedes camptorhynchus Thoms, is a salt-
marsh species in southern Australia, where it replaces A. vigilax, and
Lee has recorded Cut ex annulirostris Sk., usually a typical fresh- water
species, as breeding in brackish water.

Sandy beach.

The beach fauna of Australia is interesting, and again is largely
dipterous, although Cicindelids (C. ypsilon Dej.) are common, and various
other beetles occur in cast-up masses of seaweeds or sometimes under
rocks. The Cicindelids typify the adult insects of the beaches, for they
are pale-coloured and fast-moving, difficult to see and exceedingly difficult
to catch. The Diptera-Brachycera have similar habits. At least two
robber-flies ( Clinopogon maritima Hardy and Stichopogon minor Hardy),
two Apiocerids ( Apiocera maritima Hardy and A. pallida Norris), one
Therevid ( Anabarrhynchus maritima Hardy), one Empidid (unidentified)
and a Tabanus ( T . vetustus Walk.) frequent our beaches and, in spite of
their capacity for strong flight, seem to have a very restricted habitat.
Another pale Tabanid of the north (T. leucopterus Wulp), however, ranges
widely out to sea and has been taken on ships many miles from the land.
The Tabanidae, There vidae and Apioceridae may be classed as truly
littoral, for Miss English (1947) has recently discovered their larvae and
pupae in the sand between tide-marks, and has given a full description
of the early stages of Apiocera maritima A All these larvae are predatory,
but they obviously cannot live exclusively on each other (though they
will do so if given the opportunity), and one imagines that their major
sources of food must be the Annelids and small Molluscs which are
common beneath the surface in the same situation.

An interesting beach-fly described from Hawaii by Williams (1938)
is the greyish Dolichopodid, Asyndetus carcinophilus Par., which mounts
guard at the entrance to the burrows of the sand-crab, Oxypode ; its
larvae live in the sand and are predacious.

Nearer the sea, in fact running at the very edge of the wash, are
the smaller but equally agile flies of the Muscid sub-family Lispinae.
They are not restricted to the open beach, but have a predeliction also
for seaweed masses and the margins of pools, both salt and fresh.
Acalyptrate Diptera are also associated with seaweed, living mainly in
the decaying material at or above high tide mark. These, with
Sarcophaga and Carabid and Staphylinid beetles, constitute the “ jetsam
fauna.” The Plymouth workers list seven species, and doubtless as
many occur in Australia ; at least Phycodromiidae, Ephydridae and

Also (Proc. Linn. Soc. N.S. Wales, 74 ; 153, 1949) of Tabanus orarius Eng.

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND

Sciomyzidae are known here. Finally, one must mention Womersley’s
(1937a) discovery of the males of Pontomyia cottoni Worn. (Ghironomidae)
in small depressions at the edge of the water on a beach in South Aus-
tralia. He has also taken them with a light around jetty piles, and
thinks that they probably skim on the surface of the water (personal
communication). Those on the beaches may have been washed up with
the waves. The same species has been taken in Gunnamatta Bay, New
South Wales (Lee, 1946). We shall have more to say about this remark-
able genus later.

Rocky coast.

This is the richest in species and the best collected of the littoral
environments ; it may be taken to include the rocky shores and reefs of
sheltered waters, as well as those exposed to the open ocean. Dakin
et al. (1948) have made a detailed study of the zonation of the latter in
New South Wales, and their findings may be tabulated as a basis for
marking the distribution of the insects, although most of the records to
be considered will not be Australian, and the indications can only be
rough approximations (Table 2).

Collembola live both on the surface of supra-littoral pools and in
crannies among the rocks between tide marks. When the tide -rises, they
bury themselves in the sand ; they appear to be completely unwettable.
Womersley (1936b) described Isotoma pritchardi Worn, from the edge of
a reef in South Australia.

TABLE 2.

Ecological Zones of Rocky Coasts.

Tide

Belt

Zone

Insects*

High-tide

Supra-littoral

V. Littorinid

Thysanura (Allow achilus)
.Collembola
Coleoptera (various)
Chironomidae, Culicidae

Littoral

IV. Barnacle
III. Galeolaria

Collembola

Hemiptera (Aepophilus)
Carabidae, Staphylinidae
Parasitic Hymenoptera
Trichoptera ( Philanisus )
Tipulidae, Chironomidae,
Dolichopodidae
Spiders (Desis) and mites
(Pontarachnidae)

Zero low tide ...

Littoral-sub-

littoral

fringe

II. Pyura
I. Kelp

Chironomidae

Halobates

* Arranged systematically, not in sequence of occurrence.

There are few Hemiptera. Aepophilus, a small bug with a family to
itself near the Gerridae, is found in Europe in company with Carabid
beetles under stones and in fissures in rock not far from low water
(Miall, 1903). Species of Halobates occur in the lower zones as well as
at sea, H. whiteleggei Sk. being common in Sydney Harbour (Skuse, 1891).

Coleoptera are relatively numerous, to judge by the Plymouth
Report. Of 116 species listed from coastal Devon, 69 were classed as
coastal,” 40 as “ feub-maritime,” and 7 as “ maritime,” the last two

MARINE INSECTS

groups including representatives of seven Sub-orders (10 species of
Geodephaga, 4 Palpicornia, 23 Brachelytra, 4 Clavicornia, 1 Lamellicornia,
3 Rhynchophora, and 2 Heteromera). The notes under the species
suggest that “ sub-maritime ” corresponds with zones IV and V, and
“ maritime ” with zones I-III of the Table. Mostly the beetles occurred
under stones and seaweed or among barnacles, and only Carabids and
Staphylinids appear to extend far into the lower zones. Miall (p. 375)
has an interesting note on Aepus, one of the Carabidae : “ They run about
on stones, seaweed, sponges, etc., at low water . . . they cannot avoid
the rising tide. As soon as it reaches them, they creep under stones
and remain motionless. The body is flattened, and covered in every
part with hairs which entangle air (Audouin). There is a large pair of
air-sacs in the abdomen . . . which are no doubt useful during prolonged
submersion.” Tillyard (1926) notes three maritime Staphylinids
[Staphylinus huttoni Br., Cafius littoreus Br. and C. maritimus Br.) as
occurring in New Zealand. Though numerous in species and showing
some special adaptations, the beetles can hardly be regarded as more
than tentative intruders from the land.

Even Hymenoptera occur, Miall noting a small Proctotrupid as
having been found under stones in company with marine Crustacea on
the coast of France. Its host was not known.

There are also a few Trichoptera, the best known being Philanisus
plebejus Walk, from Australia and New Zealand. “ Its larva feeds on
coralline seaweed in rock-pools between tide-marks, and appears to be
generally distributed round the coasts of both countries. The sub-
cylindrical case is cunningly contrived from small pieces of the food-plant
and other objects, so that the larva is most difficult to detect. The imago
frequents rocky coasts and is very active.” (Tillyard, p. 394.)

Five families of Diptera are represented in addition to the jetsam
fauna mentioned above. The Culicidae are restricted to the supra-littoral
zone, where they breed in rock-pools containing various concentrations
of salt water. In New South Wales, Aedes alboannulatus Macq. only
occurs when the salt content is low (0.2-0. 7%) and Anopheles annulipes
Walk, is occasionally taken in water containing up to 1.6% salt, but
Aedes concolor Tayl. is specially adapted to these conditions, and its
larvae have been found in water with a saline content from 0.1 to 7.4%
(Woodhill, 1936). The adults are also restricted to this zone, the females
biting freely at dusk while the males hover overhead in a pre-nuptial
dance. A remarkable aberrant Culicine, Opifex fuscus Hutton, occupies
a similar niche in New Zealand (Miller, 1922) and a Ceratopogonid
(Dasyhelia) in Hawaii (Williams, 1944).

The Tipulidae occur lower in the series, their larvae being found
“ among algae on sea-rocks, submerged by the tide ” (Alexander,
1931). Almost all belong to four subgenera of the great genus
Linionia, and their adaptations seem to parallel to some extent those of
the Chironomidae (Tokunaga, 1933). Adult Dolichopodidae live in the
same zone, haunting the surf and breakers in search of their prey, and
even flying over reefs at low tide 100 yards from the shore (Miall,
Plymouth Report).

The Chironomidae are the most interesting group in this environ-
ment, and show the most complete series of progressive adaptations to a
marine existence. Stuart (1942) has described the supra-littoral species
of Scotland. The Chironominae live in, and the Tanypodinae on, the

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND

mud in brackish pools apparently similar to those described above for
the Culicines in this country. They possess haemoglobin and reduced
tracheal trunks. The Orthocladiinae occupy pools of varying salinity in
the same zone, but extend also into the upper littoral. They lack
haemoglobin, have large tracheal trunks, feed on algae, and some species
are adapted for quick emergence and oviposition ; one secretes a him
of air in the pupal case like Simuliidae. Edwards (1926) follows this
group further into the inter-tidal belt, where the larvae live among
seaweed and the adults congregate on the wet rocks. In C amptocladius
thalassophilus Goet., the females do not rise into the pre-nuptial swarm,
but wait' on the rocks for the males to descend to them. Skuse (1889)
has described C. crassipennis Sk., apparently from similar situations, in
Sydney Harbour.

In the Clunioninae, the larvae occur in the intertidal zone, sometimes
deep in pure salt water at the outer fringe, sometimes near the mouths
of streams, where there is some admixture , with fresh water ; they live
in, and mostly feed on, various species of Algae. Emergence of the
adults takes place when the tide is out. They mature quickly, and are
usually active after dark, scampering half-running, half-flying, over the
wet rocks exposed at low tide, sometimes rising in the air over the sweep
of a wave (Tokunaga, 1935), sometimes clinging submerged to the rock.
They mate on the rocks, and appear to be unwettable.

These species show progressive reduction of adult structures.
Telmatogeton and Thalassomyia have well-developed wings in both sexes,
but the males have lost the antennal plumosity, possibly associated with
loss of aerial mating. In Halirytus and Eretmoptera, there is more or
less reduction of the wings. In Clunio, the wings of the male are short
and rounded but functional, while the females have lost wings and
halteres, and are carried round attached to the males like miniature
Thynnid wasps. Recent revisions of these genera have been published
by Wirth (1947a, Thalassomyia ; 1947b, Telmatogeton) and Stone and
Wirth (1947, Clunio). Womersley (1936a) has described Telmatogeton
austr aliens Worn, from South Australia and given an account of its
biology, while Dakin et al. record Clunio pacificus Edw. from the littoral-
sublittoral fringe on the New South Wales coast.

The final step in adaptation to marine life is reached in the genus
Pontomyia, which properly belongs to the next section. Its larvae and
pupae live in delicate mud tubes among the fronds of Halophila ; the
females lack antennae, mouth-parts, wings, halteres, and all but the
stumps of the mid and hind legs ; they probably remain in the tubes
where they emerged ; while the males have reduced, distorted wings, and
swim actively in the plankton beneath the surface.

It is not to be inferred that the steps described consecutively here
represent a single line of evolution. There were three, possibly more,
lines, represented by the three subfamilies. The Orthocladiinae probably
came down via the supra-littoral pools ; the Clunioninae may have
entered the turbulence of the sea from the turbulence of rapids and
waterfalls, though Wirth suggests that the fresh-water species of
Telmatogeton are derived from marine forms ; while Pontomyia represents
an entirely different line of evolution, being derived, according to
Edwards, from the Chironomine genus T any tarsus with which it is
associated in the Halophila.

MARINE INSECTS

A brief note on marine Arachnida in Australia may be given to
complete the account of this region. Dakin et al. record the spider,
Desis crosslandi Poc., as building its webs in the Galeolaria zone, and
note its occurrence from Queensland to Victoria. We have seen what
is probably the same species on the outer part of coral reefs, nesting in
crevices inside the Lithothamnion platform at Hayman Id., and Heron
Id., Queensland. Womersley (1937b), records two Hydrachnoid mites
(Pontarachna halei Worn, and Litarachna denhami Loh.) from the littoral
zone in South and Western Australia respectively, and has also found
Halacharidae among seaweed (personal communication).

Coral reef.

The fauna of this region is an impoverished outlier of the littoral
fauna described above, its interest lying in its frequently wide detach-
ment from the land and the purity of the sea-water as indicated by the
presence of living coral. I have references to only three groups of
insects and a spider.

Collembola (Axelsonia littoralis Monz. and Pseudachorutes sp.*) occur
in the outer parts of the reef inside the rampart in similar situations to
Desis. They were not uncommon at Heron Id. The marine bug,
Halobates, lived in the same zone, and seemed more inclined to hide in
the coral than to skate on the water. In the Chironomidae, Edwards
described four species from Samoa, Clunio pacificus Edw., T any tarsus
halophilae Edw., T. maritimus Edw., and Pontomyia natans Edw., and
additional Clunioninae have been recorded from other Pacific islands.
The last three of Edwards’ species are particularly interesting, as they
represent the only truly marine Chironomine genera known, and they
were associated with Halophila, which is one of the few higher plants
to invade the sea (Buxton) .

PELAGIC FAUNA.

We have followed an ever decreasing insect fauna from the shore
to the verge of the littoral belt and to the coral reef. Now we come to
the last and smallest group. Only two kinds of insects can be described
as pelagic, the Halobatinae and the Chironomine genus Pontomyia.

Halobates and related genera skate on the surface of the oceans,
often far from land, feeding on animal remains, and laying their eggs
on floating detritus. They are unwet table, and live on, not in, the water.

Pontomyia was discovered by Buxton at Samoa. The early stages
and the female belong to the reef fauna, but the males are as truly
pelagic as the zooplankton among which they live. Buxton collected
them with a tow-net, at night, at half to low tide by sleeping the water
over the patches of Halophila. Wassell (1948) recently collected swarms
of males in a night plankton haul in 8 feet of water half a mile from the
shore in Princess Charlotte Bay, North Queensland. They were greatly
attracted by the strong light which was used to concentrate the plankton,
and swam about rapidly beneath the surface, agitating the water as the
light was moved. Many clustered on the side of the vessel at the water-
line where the light shone strongest. One could hardly imagine more
remarkable behaviour in an insect.

* Determined by Mr. H. Womersley of the South Australian Museum, Adelaide.

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND

CONCLUSION.

This review is admittedly incomplete ; references have been difficult
to trace, and some of the literature has not been available in Queensland.
Nevertheless, enough has been said to show that the sea is not as devoid
of insects as might have been imagined. Naturally, most of the species
are littoral in distribution, and the rocky coasts and reefs have proved
particularly favourable fob colonization. Among their inhabitants, there
are some which can enter the sea simply because they are unwettable
and the salt water cannot reach their bodies, but others are truly marine.
Only about ten families of half a dozen Orders have survived in the full
3.2% salinity of the open ocean, and of these the Chironomidae have
shown the greatest variety and perfection of adaptation and the strongest
tendency to populate the deeper waters. A wide field of investigation
is open to the Australian worker, for our marine insects are still but
little known, our coasts and reefs promise to be prolific collecting grounds,
and there is no reason why the fascinating study of shore ecology should
remain the exclusive province of the marine biologist.

REFERENCES.

Alexander, C. P., 1931. -The early stages of crane-flies (Diptera). Victorian
Nat.. 47 : 195-203.

Buxton, P. A., 1926. — On the colonization of the sea by insects : with an account
of the habits of Pontomyia, the only known submarine insect. ; 1 Proc . zooi.
Soc. Lond., 1926 : 807-814.

Dakin, W. J., Bennett, I., and Pope, E., 1948. — A study of certain aspects of
the ecology of the inter-tidal zone of the New South Wales coast. Aust.
J. sci. Res. (B), 1 : 176-230.

Edwards, F. W., 1926. — On marine Chironomidae (Diptera) ; with descriptions of
a new genus and four new species from Samoa. Proc. zool. Soc. Lond.,
1926 : 779-806.

English, K. M. I.. 1947. — Notes on the morphology and biology of Apiocera
maritima Hardy (Diptera, Apioceridae) . Proc. Linn. /Soc. N.S. Wales,
71 : 296-302.

Hardy, G. H., 1935. — Miscellaneous notes On Australian Diptera. III. Proc.
Linn. Soc. N.S. Wales, 60 : 248-256.

Lee, D. J., 1944. — An atlas of the mosquito larvae of the Australasian Region.
Tribes — Megarhinini and Culicini. H.Q., A.M.F. Publn., 119 pp.

Lee, D. J., 1946. — Notes and exhibits. Proc. Linn. Soc. N.S. Wales, 71: xxvi.

Lee, D. J., and Woodhill, A. R., 1944. — The Anopheline mosquitoes of the
Australasian Region. Univ. Sydney, Dept. Zool., Monograph No. 2,
209 pp.

Lindberg, H., 1937. — Okologische Studien fiber die Coleopteren und Hemipteren-
fauna im Meere in der Poio-Wiek und im Scha renarchipel von Ekenas in
Siidfinnland. Acta Soc. Fauna Flora fenn., 60 : 516-572.

Marks, E. N., 1947. — Exhibit. Larvae and adults of Culicoid.es spp. Ent. Soc.
Queensl., 14 July, 1947, Minutes pp. 3-4.

Miall, L. C., 1903. — The natural history of aquatic insects. London, Macmillan
(4th reprint, 1934) : 370-381.

Miller, D., 1922. — A remarkable mosquito., Opifex fuscus Hutton. Bull. ent.
Res., 13 : 115-126.

Skuse, F. A. A., 1889. — Diptera of Australia. Part VI. The Chironomidae.
Proc. Linn. Soc. N.S. Wales, 4 : 215-311.

Skuse, F. A. A., 1891. — Description of a new pelagic Hemipteron from Port
Jackson. Rec. Aust. Mus., 1 : 174-177.

Stone, A., and Wirth, W. W., 1947. — On the marine midges of the genus Clunio
Haliday (Diptera, Tendipedidae). Proc. ent. Soc. Washington, 49 : 201-224.

MARINE INSECTS

Stuart, T. A., 1942. — Chironomid larvae of the Millport shore pools. Trans. R.
Soc. Edin., 60 : 475-502.

Tiegs, O. W., 1949. — The problem of the origin of insects. Presidential Address
to Section D. Aust. N.Z. Ass. Adv. Sci., Hobart, 13 Jan., 1949.

Tillyard, R. J., 1926. — The insects of Australia and New Zealand. Sydney,
Angus & Robertson, 560 pp.

Tillyard, R. J., 1930. — The evolution of the Class Insecta. Pap. Proc. R. Soc.‘
Tasm. for 1930 : 1-89.

Tokunaga, M., 1933. — A marine crane-fly, Limonia ( Dicranomyia ) trifilamentosa,
of the Pacific coast of Japan, with special reference to the ecology and
the morphology of its immature stages. Philippine J. Sci., 50 : 327-344.

Tokunaga, M., 1935. — Chironomidae from Japan (Diptera). IV. The early stages
of a marine midge, Telmatogeton japonicus Tokunaga. Philippine J.
Sci., 57 : 491-511.

Wassell, J. L. H., 1948. — Exhibit. Marine insects. Ent. Soc. Queens!., 13 Sep.,
1948, Minutes p. 2.

Williams, F. X., 1936. — Biological studies in Hawaiian water-loving insects.

Part I. Coleoptera or beetles. Part II. Odonata or dragonflies. Proc.
Hawaiian ent. Soc., 9 : 235-349.

Williams, F. X., 1938. — Asyndetus carcinophilus Parent (Diptera, Dolichopodidae) .
Proc. Hawaiian ent. Soc., 10 : 126-129.

Williams, F. X., 1944. — Biological studies in Hawaiian water-loving insects.

Part III. Diptera or flies. D. Culicidae, Chironomidae, and Cera-
topogonidae. Proc. Hawaiian ent. Soc., 12 : 149-180.

Wirth, W. W., 1947a. — Notes on the genus Thalassomyia Schiner, with descrip-
tions of two new species (Diptera Tendipedidae). Proc. Hawaiian ent.
Soc., 13 : 117-139.

Wirth, W. W., 1947b. — A review of the genus Telmatogeton Schiner, with descrip-
tions of three new Hawaiian species (Diptera, Tendipedidae). Proc.
Hawaiian ent. Soc., 13 : 143-191.

Womersley, H., 1936a. — An interesting Chironomid Telmatogeton australicus
sp. n. from a South Australian reef. Rec. S. Aust. Mus., 5 : 439-443.

Womersley, H., 1936b. — Further records and descriptions of Australian Collembola.
Rec. S. Aust. Mus., 5 : 475-485.

Womersley, H., 1937a. — A new marine Chironomid from South Australia. Trans.
Proc. R. Soc. S. Aust., 61 : 102-103.

Womersley, H., 1937b. — A new species of marine Hydrachnellae from South
Australia. Trans. Proc. R. Soc. S. Aust., 61 : 173-174.

Woodhill, A. R., 1936. — Observations and experiments on Aede's concolor Tayl.
(Dipt. Culic.). Bull. ent. Res., 27 : 633-648.

— , 1931. — Plymouth .Marine Fauna. Marine Biol. Ass., Plymouth,
2nd Ed., 371 pp. Insects, pp. 223-235.

Vol. LXI, No. 4.

A NEW ERGOT FROM QUEENSLAND

By R. F. N. Langdon, M.Agr.Sc., Department of Botany,
University of Queensland.

(Received 21th June, 1949 ; read before the Royal Society of Queensland ,
31s£ October, 1949 ; issued separately ).

In 1941 an ergot or Hyparrhenia filipendula (Hochst.) Stapf was
found a few miles north of Ipswich, Queensland, but attempts to deter-
mine the species of Claviceps responsible were not successful (Langdon
1942A). In May7 1948, sclerotia were collected from this host at
Conandale, South Queensland. Subsequent germination showed that the
ergot was a species previously unknown. It was first brought to notice
by the development of the saprophyte Cerebella on infected spikelets.
The amount of honey-dew produced is limited, and after mid-morning
it usually dries up, at least on the exterior of infected spikelets, leaving
a white encrustation about the margins of the glumes. The sclerotia
remain hidden within the glumes and can be detected only by the darker
and plumper condition of the spikelets. The name of this new species of
Claviceps is derived from the unobtrusive symptom-picture shown by
infected plants.

Claviceps inconspicua Langdon ; species nova, afhnis C. annulatae
Langdon, sed stromatis colore et indumento differt.

Sclerotia fuliginea, subcylindrica vel fusoidea, in spiculis inclusa,
2-5 mm. longa. Stromata in quoque sclerotio 1 vel plures. Stipites
1.5-9 mm. longi, colore Anthracene Purple (Ridgway) vocato. Capitula
globosa, papillosa, in superhcie hyphis raris brevibus, in basi annulo
hypharum brevissimarum albarum praedita, colore Raisin Black
(Ridgway) vocato, 0.3-0. 6 mm. diam. Perithecia 155-180 X 105-125 /x.
subglobosa. Asci cylindrici, 140-175 x 4 /jl. Ascospori line ares, hyalini,
Conidia hyalina, guttulata vel granulosa, lateribus recta vel leniter curva,
hnibus ambobus rotundata, 15-20 X 5-10 /x.

In ovariis Hyparrheniae filipend.ulae (Hochst.) Stapf, Queensland.
Prope Conandale, 30th May, 1948, Langdon (425 TYPE) ; prope Ipswich,
28th May, 1941, Langdon (163) ; Grovely, Brisbane, 12th April, 1949 (426).

The sclerotia were kept dry during the winter and subjected to cold
treatment (2-4° C. for 28 days). In September they were placed on moist
sand in petri dishes. Development of the ascal stage began in mid-
November. Germination of the sclerotium begins with the protrusion
of a small white papilla which quickly grows out into a globose tuft of
white hyphae. The developing stroma is veiled with white hyphae as
it pushes up, and at maturity loose hyphal elements persist on the
surface of the capitulum. A ring of very short white hyphae is present
at the base of the capitulum where it joins the stipe. The tuft of hyphae
at the base of the stipe is persistent.

OTHER RECORDS OF ERGOT ON HYPARRHENIA.

Goncalves (1937) reported the occurrence of ergot on Hyparrhenia
rufa in Brazil, and noted an association of Cerebella with the sphacelial
stage of the disease. There are records of Cerebella and Fusarium as
saprophytes in the honey-dew of ergot of Hyparrhenia ruprechtii in
Southern Rhodesia (Hopkins, 1947). McDonald (1927) reported the

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND

occurrence of C'erebella on Hyparrhenia collina in Kenya. In Sierra
Leone, Deighton (1947) has found Cerebella on Hyparrhenia gracilescens,
H. rufa and H. subplumosa, but he stated that he did not find it associated
with ergot honey-dew. From the work of Langdon (1942B) there seems
little doubt that a record of Cerebella on a grass is a safe indication of a
prior infection of the host with ergot.

DISCUSSION.

In Queensland a number of native grasses are hosts for indigenous
species of Claviceps (Langdon 1942A). A wide search in south-eastern
Queensland since 1946 has revealed only two new hosts for ergot, and
both of these were infected by Claviceps pusilla. Hyparrhenia filipendula
is the only host known for Claviceps inconspicua. There are at present
no other suspected hosts of this ergot, i.e., plants which have been found
infected with ergot, the perfect stage of which has not been observed.
A consideration of the origin of Claviceps inconspicua, whether the fungus
is indigenous to Australia or, has been introduced, must take into account
the origin of its host. Hyparrhenia filipendula is a plant about which
there has been some doubt as to its natural distribution. Stapf (1934)
for the genus Hyparrhenia writes : “ Species over 60, almost confined to
tropical Africa (including the islands) and subtropical South Africa,
three of them extending to tropical America, one to Asia and Australia,
one to Mediterranean countries and temperate Africa.” For Hyparrhenia
filipendula, Stapf gives the general extra-African distribution as Ceylon,
the Philippines, and Australia, but adds that “ Hyparrhenia filipendula
is often found on abandoned plantations, and its occurrence in India,
Malaya and Australia may possibly be due to casual introduction.”

In Australia, Hyparrhenia filipendula is found as a constituent of
the herbage in open forest areas, and it occurs also in induced grassland
communities following changes effected by man in the natural plant cover.
Blake (1942) found Hyparrhenia filipendula associated with a number
of native grasses in an Open Eucalyptus Forest community at Running
Creek in south-eastern Queensland. The herbaceous cover was dominated
by kangaroo grass, Themeda australis, a species which is amongst the
earliest to disappear under grazing conditions. This occurrence of
Hyparrhenia filipendula in a mixture of native grasses in what must be
regarded as an area carrying almost unaltered natural vegetation is
\vorthy of note. That Hyparrhenia filipendula occurs in induced grass-
land communities is not evidence that it is an introduced grass as might
be inferred from Stapf s remarks on the occurrence of the species in
abandoned plantations. Native grasses frequently are dominant in
disturbed ground, for example, C apillipedium spicigerum and Bothriochloa
decipiens. Imperata cylindrica var. major, a species indigenous to Aus-
tralia and south-east Asia, often occupies cultivated land which has
been abandoned.

Through the courtesy of Mr. S. T. Blake of the Queensland
Herbarium, records of the locality and date of collection of specimens
of Hyparrhenia filipendula in various Australian herbaria have been
obtained. The earliest collection was by Leichhardt in 1843, the locality
being given as “ Eastern Australia.” Other early collections are from
the islands of Moreton Bay by Mueller in 1855, from the country west
of Rockhampton by Bowman in 1867, from the Clarence River (N.S.W.)
by Beckler between 1870 and 1880, and from the Apsley River in the

A NEW ERGOT FROM QUEENSLAND

Kimberley district of Western Australia by Crawford in 1887.
The distribution of Hyparrhenia filipendula in Australia, as indicated
by specimens in various herbaria, is from the Clarence River
in northern New South Wales to North Queensland, and in the
north of Western Australia. Mount Fraser, near Mossman, is the
northernmost area from which the species has been collected in
Queensland, and the grass has been recorded from a number
of coastal and sub-coastal areas at various places between its
known southern and northern limits. Crawford’s collection from the
Apsley River is the only record of the grass in Western Australia. The
comparatively late collection of Hyparrhenia filipendula in Australia
might suggest that the grass has been introduced after colonization of
Australia by white men, though its occurrence in places remote from
centres of early settlement controverts this idea. If introduced from
Africa early in the nineteenth century, the grass might be expected to
occur in the south-west of the continent or near Sydney, but it does not.
That the climatic conditions in the latitude of Sydney are such that
Hyparrhenia cannot develop to maturity there is not a tenable hypothesis.
There is in the Queensland Herbarium a fertile specimen of Hyparrhenia
(? rufa), grown in the Sydney Botanic Gardens from seed imported from
Nairobi. Although a species other than H. filipendula is concerned here,
the range of the latter in Africa does cover the territory from which the
Sydney grass was obtained. Hyparrhenia filipendula, if it had been
introduced in the Sydney or Perth areas, might have established itself
there in waste areas where it would be free from competition from native
plants. A final point against the possibility of introduction of the grass
from Africa is that Hyparrhenia filipendula does not occur south of
latitude 30° S., and so is not likely to have been brought over by travellers
who visited the Cape of Good Hope area on their way to Australia in
the late eighteenth or early nineteenth centuries. Since Hyparrhenia
filipendula was not found in the areas serving as bases for those who
originally explored or settled in other parts of the continent, it is very
likely that the record of the grass by Leichhardt in “ Eastern Australia ”
represents the collection of a naturally occurring species. The possibility
of the introduction of Hyparrhenia filipendula from south-east Asia
direct to the settlement around Moreton Bay prior to 1843 is remote.

In south-eastern Queensland, a smut, Ustilago hyparrheniae Hopkins
is common on Hyparrhenia filipendula. This smut was described from
the same host from Southern Rhodesia, and a Queensland specimen sent
to Southern Rhodesia was reported as being identical with the type
collection (Bates 1948). If seed of this grass had been accidentally
introduced to this country from Africa in the past, one might reasonably
suppose that the smut had come with it.

The occurrence of ergot on Hyparrhenia in Africa has been noted
above, but the species of Claviceps affecting the genus there has not
yet been determined. Nor is the species of Claviceps affecting
Hyparrhenia in South America known. While these ergots are undeter-
mined, one cannot say whether all the ergot diseases of Hyparrhenia are
the same. But to assume that the Australian ergot has been brought here
with an accidental introduction of seed would suppose a rather unlikely
series of events, the carriage of sclerotia, with their subsequent develop-
ment and release of ascospores at a time when the introduced host was
flowering. This view is put forward despite the presence in Australia of
Claviceps purpurea and Claviceps paspali, neither of which is indigenous

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND

to this country. The former was introduced in the nineteenth century,
probably with cereal grain or with seed of pasture grasses (possibly in
both ways). In any case, an abundance of hosts of more than one
species would be available to any germinating sclerotia, for native grasses
as well as introduced plants are recorded as hosts of this ergot in southern
Australia. Claviceps paspali appears to have been introduced much
later. It has been widespread and very common on several species of
Paspalum in eastern Australia since the summer of 1935-36, when it was
observed for the first time. The quick development of epiphytotics of
ergot in Paspalum every year now in coastal Queensland indicates how
well local conditions suit this ergot. The first sclerotia to germinate
after their introduction (probably in 1935) had available an abundance
of Paspalum dilatatum which flowers profusely, and the initial infections
should not have been difficult to accomplish. Plants of Hyparrhenia
filipendula, unlike Paspalum dilatatum the chief host of Claviceps paspali,
are not massed in pure stands over large areas, and are not common
as weeds of waste places. Nor is any alternative host of Claviceps
inconspicua known. The possibility of infection of Hyparrhenia by
ascospores from sclerotia introduced by chance at any time is very much
less than for the hosts of Claviceps purpurea and Claviceps paspali.
Furthermore, all observed occurrences of Claviceps inconspicua on
Hyparrhenia have been light infections, indicating that environmental
conditions do not usually favour epiphytotics of this ergot, and that
infection under prevailing circumstances is relatively difficult. The
enphytotic state of this ergot disease may perhaps be regarded as the
result of a long-standing association of host and parasite in this country.

The mycological evidence bearing on the question of whether
Hyparrhenia filipendula is a native or an introduced species in Australia
is divided. The introduction of a smut with the seed is feasible, but
the probability of the introduction of an ergot specific to this host is
not easy to accept. One might postulate development of an ergot species,
specific to Hyparrhenia filipendula, from some indigenous Australian
ergot. Claviceps inconspicua, morphologically, has much in common
with Claviceps pusilla, an ergot which infects a wide range of genera
in the sub- tribe Andropogoninae, and with Claviceps annulata, an ergot
infecting Eulalia of the sub-tribe Saccharinae. Possibly Claviceps
inconspicua and Claviceps annulata are Australian variants of the more
widely distributed Claviceps pusilla. If one rejects the hypothesis that
Claviceps inconspicua is an evolutionary product of the past century,
specific to Hyparrhenia filipendula and developed since the time of that
grass’s introduction to Australia, the above proposition may still be
valid. The host Hyparrhenia filipendula is known from the Philippines
(Merrill 1925) and from Ceylon, India and Malaya (Stapf 1934). Possibly
it is a species of wide natural distribution, extending from Africa, through
Asia, to Australia. If it is a grass of long-standing occurrence in Aus-
tralia, an explanation of the presence here of its ergot, having affinities
with other indigenous ergots, can be found.

The ecological, phytogeographical and mycological evidence pre-
sented here supports the theory that Hyparrhenia filipendula is a species
native to Australia. If one accepts the indigenous nature of the host,
then Claviceps inconspicua can be regarded as an ergot indigenous to
Australia .

A NEW ERGOT FROM QUEENSLAND

ACKNOWLEDGMENTS.

I wish to thank Mr. S. T. Blake for his assistance in the preparation
of this paper by discussion with me of the plant distribution problem
involved and by making available various records of the occurrence of
Hyparrhenia filipendula in Australia ; and Professor D. A. Herbert whose
constructive criticism of the theories put forward has been most helpful.
Financial assistance for this work was granted by the University of
Queensland Commonwealth Research Projects Committee, to whom
the author is grateful.

REFERENCES.

Bates, G. R., 1948. — Private communication.

Blake, S. T., 1942. — Queensl. Nat. 13 : 4-12.

Deighton, F. C., 1947. — Private communication.

Goncalves, R. D., 1937.— O Biologico 3 : 74-75.

Hopkins, J. C. F. , 1947. — Private communication.

Langdon, R. F. , 1942a. — Proc. Roy. Soc. Queensl. 54 : 23-32.

Lajntgdon, R. F. , 1942b. — Phytopathology 32 : 613-617.

McDonald, J., 1927. — Ann. Rept. Dept. Agric. Kenya, p. 229.

Merrill, E. D., 1925. — Enumeration of Philippine Flowering Plants, Vol. 1.
Stapf, O., 1934. — In Prain, D.; Flora of Tropical Africa, Vol. 9.

Vol. LXI., No. 5.

REVISION OF BREGMACEROS WITH
DESCRIPTIONS OF LARVAL STAGES
FROM AUSTRALASIA

By Ian S. R. Munro, M.Sc., Division of Fisheries, Commonwealth
Scientific and Industrial Research Organization.

(With Ten Figures in the Text).

( Received 25th October, 1949 ; tabled before the Royal Society of
Queensland, 28 th November, 1949 ; issued separately — — — — — — -).

SUMMARY.

Six species of the genus Bregmaceros are recognised, including
B. rarisquamosus sp. nov. from New Guinea and the Solomon Islands.
All are described, references to species listed, and the distribution of the
genus is given. On the basis of larval and post-larval stages,
B. macclellandi is recorded from eastern Australia and B. japonicus
and B. nectabanus are recorded from eastern Australia and New Guinea.
The larval and post-larval stages are described and figured.

INTRODUCTION.

A large series of plankton collections obtained off the eastern coast
of Australia during the period 1938 to 1942 by F.R.V. “ Warreen,”
fisheries research vessel of the Commonwealth Scientific and Industrial
Research Organization, has yielded seventy-two larval fishes of the genus
Bregmaceros. F.R.V. “ Stanley Fowler,” another survey vessel of this
organization, obtained by means of a submarine lamp six specimens from
Northern Territory and North-Western Australia in 1949. During 1948
to 1950, M.V. “ Fairwind,” fisheries survey vessel of the Department of
External Territories, obtained by means of a submarine lamp fourteen
additional specimens in Papua, New Guinea, and the Solomon Islands.

The identification of Australasian material has necessitated a review
of the literature dealing with all described forms. Type material has
not been accessible, but the differences between species have been obtained
from published descriptions and figures. Compilations of adult characters
and complete lists of references have been drawn up for each of the six
species recognised. A revised key has been prepared to distinguish at
least the adults of the accepted species. The distribution of the genus
has been summarised by means of a map (Text Fig. 1).

The genus Bregmaceros was proposed by Thompson (1840, p. 184)
for B. macclellandi from the Ganges River. Gunther (1889, p. 24) rightly
placed Calloptilum mirum Richardson (1843, p. 46) from China Seas and
Asthenurus atripinnis Tickell (1865, p. 32) from Burma (Akyab) in the
synonymy of B. macclellandi. Five other forms have since been described,
some of which have been considered worthy only of sub-specific or
varietal rank. These are B. atlanticus Goode and Bean (1886, p. 165)
from the West Indies, B. bathymaster Jordan and Bollman (1889, p. 173)
from the Gulf of Panama, B. longipes Garman (1899, p. 191) from western
Mexico, B. japonicus Tanaka (1908, p. 42) from Japan, and B. nectabanus
Whitley (1941, p. 25) from Darwin, northern Australia.

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND

The status of the various forms is poorly understood. Parr (1931,
p. 48) recognised the need for a revision and presented a key to distinguish
four species. On the other hand, Norman (1930, p. 339) followed the
simpler course. By ignoring differences, he united all Atlantic, Pacific
and Indian Ocean material in a single species. This view is not accepted
as larval material from eastern Australia is composed of three species,
and three species occur in adjacent localities in New Guinea. This
immature material serves to show that authors have been in error in
assuming that variations in pigmentation are due to different stages in

KEY TO SPECIES.

I. Less than 70 scales in longitudinal series :

1. Ventral fins half body length without caudal ; less than 50 scales in longi-
tudinal series ; body and fins pale and hyaline B. rarisquamosus

2. Ventral fins two-thirds body length without caudal ; more than 50 scales in
longitudinal series ; body and fins in part dusky or with numerous black dots :

A. 10 scales in transverse series :

a. Eye 3.0 in head, nearly twice snout ; interorbital less than eye ;
body silvery with rows of black dots near bases of dorsal and anal
fins B. batKy master

aa. Eye 3.5 to 4.0 in head, equal to or shorter than snout ; interorbital
conspicuously wider than eye ; body uniformly dark ... B. atlanticus

A A. 14 to 16 scales in transverse series ; body silvery, minutely dotted with
brown; at least dorsal fins black ,... B. macclellandi

II. More than 70 scales in longitudinal series :

1. 13 or 14 scales in transverse series ; eye less than interorbital and snout ;

depth more than 8 in length without caudal, less than height of anal rays ;
body dusky ; fins dark B. japonicus

2. 17 scales in transverse series ; eye greater than interorbital and snout ; depth

less than 7 in length without caudal, nearly equal to height of anal rays ;
body pale with brown dorso-lateral stripe ; fins pale B. nectabanus

REVISION OF BREGMACEROS : DESCRIPTIONS OF LARVAL STAGES 39

Bregmaceros rarisquamosus sp. nov.

(Fig. 10)

D. (11-14) -I- (7-12) + (14-18), (36-39). A. (12-15) + (7-12) +
(15-18), (38-40). P. 15-16. C. 24-26. Lat. sc. 43-50. Trans, sc. 10-12.
Depth 5.0 to 6.1, head 5.1 to 5.6 in body length without caudal. Eye
very large, 2.7 to 3.5 in head, 0.25 to 0.5 greater than snout, 1.3 to 1.7 in
interorbital. Maxilla extends to below the posterior third of the eye,
1.9 to 2.1 in head. Nuchal appendage equal to, or 0.2 greater than head.
Dorsal fin inserted slightly in advance of the anal fin. Longest anal ray
0.9 to 1.1 in head. Ventrals 0.45 to 0.5 of body length without caudal,
extending to half way along the anterior raised portion of the anal fin.
Pectorals 1.4 to 1.6 in head. (See Table I).

Eye black. Flesh glassy transparent in life, white when preserved.
Body ornamented with large stellate melanophores. An hour-glass-
shaped patch present on the postero-dorsal aspect of the head between
the eyes and the nuchal appendage consisting of five small clusters, the
three anterior composed of large melanophores and the two posterior of
small spidery melanophores. On the dorso-lateral aspect of each side
of the body is a weak, sub-horizontal stripe, extending from the upper
angle of the operculum to the caudal peduncle, composed of two or more
series of melanophores. The cells are largest under the posterior elevated
part of the dorsal fin, and smallest under the low middle section of the
dorsal fin. The vertebral column is heavily pigmented with large internal
melanophores. A single series extends along the ventral mid-line of the
trunk between the origins of the ventral and anal fins. Fins hyaline.
Rays of posterior part of dorsal fin lightly dotted with minute
melanophores.

This species is closest to B. nectabanus Whitley. Both species are
similar in coloration, but fin and scale counts differ considerably. The
New Guinea species has fewer dorsal and anal rays, and considerably
less horizontal and vertical tracts of scales. It has a larger eye and
shorter ventral fins.

Based on two sexually mature females (28.5, 33.3 millimetres) from
Bostrem Bay (Sek Harbour), north coast of New Guinea (27.12.48) and
three young adults (23.0, 24.5, 25.5 millimetres) from Port Moresby
Harbour, Papua (2.7.48). Post-larval stages were obtained at Madang
Harbour, north coast of New Guinea and Kieta Harbour, eastern
Bougainville, Solomon Islands. All were attracted to the surface at
night using a submarine lamp. The material was collected on the
M.V. “ Fan-wind.” The largest adult female from Bostrem Bay is
selected as holotype and deposited at the Marine Biological Laboratory,
Division of Fisheries, Commonwealth Scientific and Industrial Research
Organization.

Larval Stages : — Five post-larval stages are referred to this species.

Length

Locality

Date

Dorsal Fin Rays

Anal Fin Rays

16.4 mm.
17.0 mm.
18.2 mm.
19.7 mm.
215 mm.

Kieta, Bougainville
Kieta, Bougainville
Kieta, Bougainville
Kieta, Bougainville
Madang, New Guinea ...

21.10.49

22.10.49

21.10.49

22.10.49

26.11.49

13 + 12 + 14, (39)
14+ 9+15, (38)

14 + 12 + 14, (38)
13+ 7 + 18, (38)
13 + 11 + 15, (39)

14+ 8 + 17, (39)
15+ 9 + 15, (39)
12+ 8 + 18, (38)
14 + 10 + 15, (39)
12 + 10 + 18, (40)

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND

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REVISION OF BREGMACEROS ! DESCRIPTIONS OF LARVAL STAGES 41

16.4 to 21.5 millimetre post-larvae. — Most of the adult facies present.
Depth 6.0 to 7.0, head 4.5 to 5.5 in body length without caudal. Eye
relatively smaller than in adults, 4.0 to 4.5 in head, equal or 0.2 greater
than snout, 0.2 less than to 0.2 greater than interorbital. Maxilla extends
to below posterior border of pupil. Nuchal appendage equal to or 0.2
less than head, not reaching origin of dorsal hn. Dorsal fin inserted
above or slightly in advance of anal fin. Ventrals 0.4 to 0.45 of body
length without caudal. Pectorals 0.5 to 0.7 of head length. Scales
present but difficult to count ; one example from Madang has 11
transverse and 49 lateral series. Flesh white in preserved condition,
transparent in life. Pigmentation similar to adults but melanophores
are smaller and less numerous. The dorso-lateral stripe is composed
of a single series of small melanophores restricted to the caudal region.
Pigmentation is insufficient to distinguish these post-larvae from those
of B. nectabanus of similar size and development.

Bregmaceros nectabanus Whitley.

Bregmaceros nectabanus Whitley 1941, p. 25, fig. 18 (Darwin, Northern
Territory, Australia — Type locality).

Bregmaceros macclellandi (non Thompson) Kent 1889, p. 240 (Cambridge
Gulf, NW. Australia). McCulloch 1923, p. 29 (Darwin).

Paradice and Whitley 1927, pp. 81, 97 (Darwin).

D. (12-18) -f (9-16) + (17-23), (40-55). A. (15-19) + (9-12) +
(17-24), (42-53). C. 28. Lat. sc. 70-74. Trans, sc. 17-18. Depth 6.1
to 6.6, head 5.2 to 5.9 in body length without caudal. Eye small to
moderate, 3.4 to 3.8 in head, equal or 0.1 greater than snout and equal
or 0.2 greater than interorbital. Maxilla reaches to below posterior
border of eye, 1.6 to 2.0 (1.8) in head. Nuchal appendage 0.1 to 0.4
longer than head. Dorsal fin inserted slightly in advance of the anal fin.
Longest anal rays 0.8 to 0.9 of head length. Ventrals 0.5 to 0.6 of body
length without caudal, extending to end of anterior raised portion of
anal fin. Pectorals 1.4 to 1.7 in head. (See Table II).

Eye black. Flesh yellowish-white in spirits, probably transparent
in life. Body ornamented with large, indistinct stellate melanophores.
A group is present on the postero-dorsal aspect of the head. An indistinct
brownish stripe composed of one or two series of melanophores extends
along each side from the upper angle of the operculum to the caudal
peduncle. The fins are hyaline. The pigmentation is similar to that
of B . rarisquamosus but the melanophores are smaller and more numerous.

Based on the holotype (Australian Museum Reg. No. I A 1719)
collected at Darwin in 1923 by Dr. W. E. J. Paradice during survey
work by H.M.A.S. “ Geranium.” Additional adult material was obtained
by F.R.V. “ Stanley Fowler ” in Northern Territory and North-Western
Australia. This consists of four specimens from Marchinbar Island,
Wessel Group (18.10.49), one from Timor Sea, 30 miles WNW. of
Charles Point, Northern Territory (22.9.49) and one from Mission Bay,
Napier Broome Bay, Western Australia (11.12.49). All were attracted
to the surface at night by a submarine lamp. Occurrence of this species
on the north coast of New Guinea is based on three post-larval stages
from Wewak Harbour collected on 23.11.49 from M.V. “ Fairwind ”
with the aid of a submarine lamp. The distribution is also extended
to the coasts of Queensland and New South Wales, based on sixty-four

Table II. — Fin Ray Counts, Scale Counts and Body Proportions of Seven Individuals of Bregmaceros nectabanus.

42 PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND

Napier Broome
Bay

34.0 mm.

15 + 16+22, (53)
17 + 11+24, (52)
71

5.4
6.6

3.4
1.1
1.0

1.5
1.8

1.3

1.1

1.8

Timor Sea

30.0 mm.

12 + 10 + 18, (40)
16 + 10 + L7, (43)
70

5.2
6.1
3.6
1.0
1.0

1.4

2.0

1.3
1.2
2.0

Wessel Is.

30.5 mm.

16 + 12 + 17, (45)
15 + 10 + 17, (42)
70

5.5

6.2

3.8
1.0
1.0

1.4

1.8

1.2

1.1

2.2

Wessel Is.

39.5 mm.

15 + 15 + 21, (51)
17 + 11+22, (50)
73

5.5

6.5

3.8
1.0
1.0

1.5

1.8

1.4

1.1

2.0

Wessel Is.

49.8 mm.

17 + 12+22, (51)

18 + 11+24, (53)

5.5

6.5

3.7
1.1
1.0

1.5

1.8

1.3

1.2

1.8

Wessel Is.

54.0 mm.

17 + 15+23, (55)
16 + 12+22, (50)
74

5.9

6.6

3.6

1.0

1.5

1.6

1.2

1.7

Holotype

Darwin

32.0 mm.

+ 16 + 20, (50)
+ 11+23, (53)
73

5.8

6.5

3.7
1.0
1.2

1.7

1.8

1.1

1.2

1.6

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REVISION OF BREGMACEROS : DESCRIPTIONS OF LARVAL STAGES 43

post-larval stages from plankton collections made by F.R.V. £< Warreen ”
in the vicinities of Lady Elliot Island, Break-Sea Spit, Cape Moreton,
Cape Byron, Coffs Harbour, Trial Bay, Crescent Head, Crowdy Head,
Port Stephens, Sydney Heads, Jervis Bay, and Bermagui.

Larval Stages. — The immature specimens from Wewak measure
16.5, 19.0 and 19.5 millimetres respectively. The sixtv-four planktonic
specimens obtained by F.R.V. “ Warreen ” vary in length from 2.1
to 21.7 millimetres.

Mo. of
Speci-
mens

Size Range

Station

Position

Date

Net

Depth

16.8 mm.

40A/38

26°'54'S. 153° 24' E.

20.9.38

N70

25 m.

8.4-21.7 mm.

, ,

,, , ,

, ,

N200

0 m.

9.1 mm.

46/38

24° 20' S. 153° 02' E.

19.9.38

N70

25 m.

4. 2-9. 8 mm.

, ,

, , , ,

, ,

N200

0 m.

14.0 mm.

48/38

27° 02' S. 153° 45' E.

21.9.38

N100

0 m.

14.7-16.8 mm.

49/38

28° 37' S. 153° 42' E.

21.9.38

N200

0 m.

11.9 mm.

52/38

30° 16' S. 153° 32' E.

23.9.38

N100

0 m.

9.1 mm.

128/39

32° 37' S. 152° 22' E.

3.5.39

N100

25 m.

4. 2-6. 3 mm.

133/39

28° 38' S. 153° 43' E.

6.5.39

N70

25 m.

8.0-10.8 mm.

, , , ,

, ,

N100

25 m.

15.4-16.1 mm.

136/39

27° 03' S. 153° 31' E.

14.5.39

N70

25 m.

16.8 mm.

, y

,, , ,

N100

0 m.

14.7-16.8 mm.

■ a

. .

N100

25 ,m.

11.9 mm.

137/39

30° 55' S. 153° 08' E.

16.5.39

N70

25 m.

14.0 mm.

, ,

, , , , ,

, ,

N100

25 m.

16.1-19.6 mm.

139/39

31° 51' S. 152° 50' E.

17.5.39

N70

25 m.

14.0-18.9 mm.

, , , ,

, ,

N100

25 m.

8.4 mm.

196/39

24° 15' S. 153° 03' E.

7.7.39

N200

0 m.

2. 1-4.2 mm.

203/39

Off Crescent Head ...

18.7.39

N100

9-200 m.

3.9 mm.

31/40

24 miles SE. of

Sydney Heads

25,4.40

N70

0-200 m.

7.0 mm.

33/40

15 miles ENE. of

Jervis Bay

30.4.40

N70

0-200 m.

4.9 mm.

, ,

, , , ,

N100

0-200 m.

11.2 mm.

73/41

12 miles ENE. of

Bermagui

12.10.41

N100

0-50 m.

3.9 millimetre post-larva. — (Fig. 2). Yolk completely absorbed.
Mouth and intestinal tract functional. Body short relative to depth.
Head and visceral cavity disproportionately large. Eye black ; choroid
fissure incompletely closed. Maxilla extends to below middle of pupil.
39 or 40 myomeres. Fins little differentiated. Nuchal appendage
present. Ventral fins represented by rudiments divided into 3 unequal
rami. Pectoral present, consisting of an undivided fold and a muscular
base. Dorsal, anal and caudal fins represented by a continuous fin fold
in which rays of each fin are incompletely differentiated. Pigment
entirely lacking.

8.0 millimetre post-larva.— (Fig. 3). Considerable increase in develop-
ment of body form and differentiation "of fins. Body more elongate
than in 3.9 millimetre larvae. Head 4.5, depth 5.0 in body length
without caudal. Eye has lost choroid fissure ; 3.5 in head, slightly less
than snout. Maxilla extends to below pupil. Branchiostegal rays
plainly visible. Pectoral fin 0.75 of "head length. Nuchal appendage

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND

I.S.R. MUNRO Del. ^

Text Figs. 2-4. — Fig. 2: Bregmaceros nectabanus Whitley, 3.9 millimetre postlarva from “ Warreen ” station 31/40 (off Port
Hacking). Fig. 3: Bregmaceros nectabanus Whitley, 8.0 millimetre postlarva from “Warreen” station 133/39 (off Cape Byron).
Fig. 4: Bregmaceros nectabanus Whitley, 16.1 millimetre postlarva from “ Warreen ” station 136/39 (off Cape Moreton).

REVISION OF BREGMACEROS ! DESCRIPTIONS OF LARVAL STAGES 45

extends to origin of dorsal fin ; slightly exceeds head length. Ventral
fin rays (3) extend to the end of anterior part of anal fin ; reach 0.4 of
the body length without caudal. Rays completely differentiated in all
fins. Dorsal and anal separated from caudal. Caudal rounded or slightly
pointed. In specimen figured D. 43, A. 45, C. 26. Anal and dorsal
fins not differentiated into high and low parts. Eye black. Body
otherwise unpigmented. First appearance of chromatophores is at
9.0 millimetres, when a few large stellate melanophores develop on the
caudal base.

16.1 millimetre post-larva. — (Fig. 4). Typical of series which range
from 14.0 to 21.7 millimetres. Most of adult facies present. Head
4.8, depth 7.0 in body length without caudal. Eye small, 4.5 in head,
less than snout or interorbital ; equipped with an adipose lid and pig-
mented black. Maxilla extends to below posterior border of pupil.
Nuchal appendage equal to head length, does not quite reach to origin
of dorsal fin. Ventrals reach anterior tip of anal but less than 0.5 of
body length without caudal. Anal and dorsal fins with elevated anterior
and posterior sections as in adults. In specimen figured, D. 14 -f- 11 -f- 17,
(42), A. 17 + 12 + 18, (47). In the Wewak specimens D. 18 + 12 -f- 17,

(47) ; 14 + 10 + 18, (42) ; 14 + 10 + 19, (43) and A. 18 + 9 + 21,

(48) ; 16 + 10 + 18, (44) ; 16 + 10 + 17, (43). Caudal now slightly
emarginate. Pectorals 0.6 of head length, with 15 or 16 rays. Scales
present but difficult to count ; one example has 17 transverse and 70
lateral series. Body white or pinkish in preserved condition, probably
transparent in life. Several series of stellate melanophores present.
A patch of small melanophores on postero-dorsal aspect of head. An
oblique row of single series extends from angle of operculum to origin
of dorsal fin. A few large melanophores between bases of pectoral and
ventral fins. An internal cluster lines the upper surface of the visceral
cavity. Four to six large, stellate chromatophores on caudal peduncle.
A single series continues forward above the lateral mid-line to the origin
of the posterior elevated portion of the dorsal fin. Some internal
melanophores invest the vertebral column in the caudal region. All
fins hyaline.

Bregmaceros macclellandi Thompson.

Bregmaceros macclellandi Thompson (ex Cantor’s MS) 1840, p. 184, fig. 6
(Ganges Delta). Gunther 1862, p. 368 (China Sea ; Philippine
Islands). Day 1865, p. 171 (Malabar and Bengal Coasts). Day
1875-1878, p. 418 (India). Day 1889, p. 433, fig. 151 (Bombay
Coast, Burma, Andaman Islands). Gunther 1889, pp. 22-25,
pi. 3, figs. A, B (Indian Ocean, Pacific Ocean, Amboina, Indian
Archipelago). Alcock 1893, p. 181 (Bay of Bengal). Alcock 1899,
p. 75 (Bay of Bengal, Andaman Islands, Malabar Coast). Weber
1913, p. 174 (Madura Sea, Bima Bight, Molo Straits, Borneo Bank,
N. Celebes (Kwandang Bay), Molucca Passage, Halmahera Sea,
Waigeu, W. Ceram (Kawa Bay), Sula-Besi (Sanana Bay), Banda
Sea, Wowomi-Buton, Buton Straits, S. Celebes-Saleyer, Ambon,
Kei Islands, Savu Sea, N. Soembawa (Salah Bight), Flores Sea).
Gilchrist and Thompson 1914, p. 87 (Cape Natal). Gilchrist and
Thompson 1917, p. 320. Barnard 1925, p. 325 (Agulhas Bank ;
Natal). Weber and Beaufort 1929, p. 6, fig. 2 (N. Java, Samarang
Road). Smith 1933, p. 53 (Siam). Delsman and Hardenberg 1934,
p. 32, fig. 23.

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND

Calloptilum mirum Richardson 1843, p. 95, pi. 46, figs. 4-7 (China Seas).
Asthenurus atripinnis Tickell 1865, p. 32, pi. 1 (Bay of Bengal off Akyab).

Bregmaceros atripinnis Day 1869, p. 522. Day 1875-1878, p. 418, pi. 91,

fig. 1.

Bregmaceros sp. Wood-Mason and Alcock 1891, p. 29 (Bay of Bengal,

off mouth of Kistna River).

D. (15-20) +. (10-17) + (13-22), (41-57). A. (18-22) + (10-16) -f
(15-26), (43-63). Lat. sc. 54-71. Trans, sc. 13-16. Depth 5.5 to 7.0,
head 5.5 to 7.0 in body length without caudal. Eye moderate, 3.5 to
4.5 in head, equal to or slightly less than interorbital and snout. Maxilla
extends to below middle of eye, 2.1 to 2.5 in head. Nuchal appendage
0.4 to 0.5 longer than head. Dorsal fin inserted slightly in advance of
anal fin. Longest anal rays 0.25 greater than head. Ventrals 0.63 of
body length without caudal, extending past end of first section of anal
fin. Pectorals equal to head without snout. Nape and back brown.
Cheeks and flanks silvery or greenish, minutely dotted with small brown
chromatophores. Dorsal, pectoral, anal and caudal fins blackish.
Ventrals whitish. In young, fins hyaline with peripheral portions
blackish. Pharyngial and abdominal epithelia black. (Compiled).

The Australian Museum has a single example (Reg. No. B 7536)
from Bombay, 79 millimetres total length, procured from Dr. Francis
Day in 1885 as B. atripinnis. D. 20 + 19 + 23, (62). A. 20 + 16 4- 25,
(61). Lat. sc. 76. Trans, sc. 14. Head 6.5, depth 6.5 in body length
without caudal. Eye 3.0 in head, equal to interorbital, 1.4 in snout.
Maxilla extends to below posterior edge of pupil, 2.0 in head. Nuchal
appendage twice head. Dorsal fin inserted slightly in advance of anal
fin. Longest anal rays 0.25 greater than head. Ventrals 0.5 of body
length without caudal. Pectorals 0.9 of head length. Colour brownish ;
skin minutely dotted with brown specks, about 3 to 5 under each scale.
Pectoral and caudal fin dusky. Dorsal dark distally. Ventrals and
anal white. Pharyngial epithelium black.

Distributed throughout the Indo-Pacific, including eastern Africa,
India, Burma, Andaman Islands, China, Philippine Islands and Nether-
lands East Indies. Former records from Australia (Darwin and
Cambridge Gulf) refer to B. nectabanus. Although adults are unknown
from Australian seas, larvae have been obtained in plankton nets by
F.R.V. “ Warreen ” from Queensland (Break-Sea Spit) and New South
Wales (CofLs Harbour and Narooma).

Larval stages— 1 prolarva and 4 post-larvae are included in the
plankton collections and they are identified as B. macclellandi on the
basis of pigmentation and body proportions.

Specimens

Station

Position

Date

Net

Depth

11.2 mm.\
13.6 mm./

52/38

30° 16' S.

153° 32' E.

23.9.38

N100

0 m.

12.6 mm

144/39

36° 15' S.

150° 24' E.

31.5.39

N200

100 m.

6.6 mm.

195/39

24° 2L S.

153° 22' E.

7.7.39

N100

0-200 m.

5.8 mm.

30/40

30° 18' S.

153° 32' E.

22.4.40

N100

0-200 m.

5.8 millimetre prolarva. — (Fig. 5). Yolk almost completely absorbed
Mouth large, functional. Intestine of several clearly defined coils. Eye
black, choroid fissure not closed. Pectoral and ventral fin rudiments.

REVISION OF BREGMACEROS ! DESCRIPTIONS OF LARVAL STAGES

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND

present. Unpaired fins represented by a continuous fin fold. No fin
rays differentiated. Nuchal appendage either undeveloped or detached.
Ventral fins represented by a single elongate process, not differentiated
into rays. About 45 myomeres. Four dorsal and two ventral large,
stellate melanophores at junction of myomeres and fin folds. Smaller
melanophores on caudal part of fin fold, fleshy base of pectoral rudiments,
intestinal loops and supracephalic sinus. General facies of this prolarva,
especially the elongate ventral fin rudiment, indicate identity with the
genus Bregmaceros. The large melanophores which are carried over into
later stages, indicate this particular species.

6.6 millimetre post-larva. — (Fig. 6). Body short relative to depth.
Head and visceral cavity disproportionately large. Eye lacking choroid
fissure ; black. Operculum and branchiostegals clearly differentiated.
All radials of dorsal, caudal and anal clearly visible. D. 50. A. 52.
C. 26. Caudal fin rounded. Both dorsal and anal fins elevated anteriorly
and posteriorly. Nuchal appendage present but probably broken.
Ventrals divided into 3 unequal rays. A reticulum of small, stellate
melanophores invests the dorsal aspect of visceral cavity. A few are
scattered over cheeks and base of pectoral fin. Four dorsal and four
ventral giant, stellate melanophores on trunk. They are internal to the
musculature and probably represent those on the fin folds of 5.8 millimetre
larvae.

11.2 to 13.6 millimetre post-larvae'.— { Fig, 7). Advanced larvae
measuring respectively 11.2, 12.6 and 13.6 millimetres, agree closely in
all characters and appear to be later stages of the 5.8 and 6.6 millimetre
larvae described above. Body form more closely approaches that of
adult B. macclellandi . Head 4.5, depth 5.75 in body length without
caudal. Eye 4.0 in head, equal to snout, slightly less than interorbital.
Maxilla extends to slightly behind centre of eye. D. 47-48. A. 10 +
22 -f 17, (49). C. 30, slightly emarginate. Ventrals 0.5 of body length
without caudal. Pectorals 0.6 of head length ; with 15 rays. Scales
developed in largest specimen ; 14 transverse series ; lateral series
indeterminate. Body pigmented with numerous small, stellate melano-
phores as noted by previous authors in the young of B. macclellandi.
They are larger and arranged differently from those of B. japonicus
larvae. In the region of the anterior parts of dorsal and anal fins are
5 or 6 longitudinal rows. Under the posterior part of the dorsal fin
and on caudal peduncle are 7 or 9 such rows. Others are present on
nape, cheeks, lips, breast, belly and fleshy base of pectoral. Unpaired
fins heavily pigmented, especially the posterior parts of dorsal and anal
and caudal base. Fin pigmentation consists of series of elongate
melanophores distributed along the fin rays. They are packed closehT
together and partly cover membranes of posterior parts of dorsal and
anal fins. Larvae of this species are shorter and greater in cross-section
than larval B. japonicus of similar size and development.

Bregmaceros japonicus Tanaka.

Bregmaceros atlanticus japonicus Tanaka 1908, p. 42, fig. — (Sagami

Sea, Japan — Type locality). Parr 1931, p. 49.

Bregmaceros japonicus Tanaka 1913, p. ISO, pi. 51, fig. 197 (Sagami Se?,

Toyama Bay, Kagoshima). Jordan, Tanaka and Snyder 1913,

p. 406. Tanaka 1933, p. 332 and fig. — . Okada 1938, p. 270.

REVISION OF BREGMACEROS : DESCRIPTIONS OF LARVAL STAGES 49

D. (15-17) + 20 + (20-23), (55-60). A. (23-32) + (2-6) -f (23-24),
(52-58). Lat. sc. 72-75. Trans, sc. 13-14. Depth 8.5 to 8.6, head
6.8 to 6.9 in body length without caudal. Eye 3.3 to 5.0 in head, less
than interorbital and snout. Maxilla extends to posterior border of
pupil, 2.3 in head. Nuchal appendage 0.6 longer than head. Dorsal
tin inserted directly above anal tin. Longest anal ray 0.5 longer than
head. Ventrals 0.6 of body length without caudal, extending almost to
end of low part of anal fin. Pectorals equal to distance from centre
of pupil to posterior end of head. Body dusky1 ; back very dark. Dorsal,
caudal and pectoral fins dark. Ventral and anal tins dusky. Inner
lining of operculum black. (Compiled).

Hitherto known only from Japan. Although adults are unknown
from Australian seas, planktonic larvae have been obtained by F.R.V.
“ Warreen ’’ from off the coast of New South Wales (Coff’s Harbour,
Crescent Head and Port Hacking). The species is known also from
northern New Guinea on the basis of a 25.0 millimetre specimen from
Madang Harbour, collected by M.V. “ Fairwind ” (26.10.49) using a
submarine lamp. Considered by some to be a form of B. atlanticus.

Larval stages.— There are 3 post-larvae which differ from those of
B. macclellandi in pigmentation and proportions. Their elongate bodies
and more numerous fin rays identify them as B. japonicus.

Specimens

Station

Position

Date

Net

Depth

1 1.5 mm.

50/38

28° 37' S. 153° 54' E.

22.9.38

N200

0 m.

21.0 mm.

104/38

34° 3' 30" S.15 1° 39' E.

15.12.38

N200

0 m.

22.4 mm.

203/39

Off Crescent Head

18.7.39

N100

0-200 m.

11.5 millimetre post-larva. — (Fig. 8). Development is slightly less
advanced than in the largest post-larva of B. macclellandi, from which
it differs in having a more elongate body, greater numbers of dorsal and
anal fin rays, and a pigmentation of smaller and more numerous
melanophores. Head 5.5, depth 7.5 in body length without caudal.
Eye equal to snout, 4.0 in head. Dorsal fin inserted slightly behind anal.
Caudal fin rounded, whereas larval B. macclellandi of equal length has
emarginate fin as in adult. Ventrals less than 3.0 in body length without
caudal. D. 52-53. A. 17 -f- 12 -f- 28, (57). Numerous small stellate
melanophores are scattered over the entire body and comprise 8 to 10
irregular horizontal rows. Visceral region is unpigmented except for a
mid- ventral series of melanophores between the breast and anus. Others
are present on supracephalic sinus, snout, preoperculum and mandible.
Unpaired fins hyaline except for a few small melanophores on the basal
parts of the posterior rays of dorsal and anal, and a few scattered on
the caudal rays.

21.0 to 22.4 millimetre post-larvae. — (Fig. 9). Form and proportions
are similar to adults of B. japonicus. Head 5.6, depth 7.0 to 9.0 in
body length without caudal. Eye 1.0 to 1.5 in snout and interorbital,
5.0 in head, black and equipped with an adipose lid. D. 14 -J- 16 -f- 23,
(53). A. 20 + 11 + 25, (56). Dorsal fin inserted noticeably behind
anal origin. Ventrals extend 0.36 to 0.5 of the body, length without
the caudal. Pectorals 2.5 in head, with 16 to 17 rays. Maxilla reaches
almost to hind border of eye. Nuchal appendage short but may be
broken. Body pigmented with numerous, small, stellate melanophores

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND

I S. R. MUNRO Del

Text Figs. 9 and 10. — Fig. 9 : Bregmaceros japonicus Tanaka, 21.0 millimetre postlarva from “ Warreen ” station 104/38
(off Sydney Harbour). Fig. 10 : Bregmaceros rarisquamosus sp. nov., 33.3 millimetre female from Bostrem Bay, Sek Harbour, North Coast

of New Guinea. Scale tracts not shown.

REVISION OF BREGMACEROS ! DESCRIPTIONS OF LARVAL STAGES 51

scattered irregularly over most of head and trunk. Cheeks and posterior
part of visceral region unpigmented. Interorbital region with a pro-
minent cluster of melanophores. A few small melanophores present on
lips and fleshy bases of pectoral and ventral fins. Dorsal, anal and
caudal fins hyaline, distinct from the heavily pigmented unpaired fins
of B. macclellandi . At most, a few melanophores on the basal parts of
the anterior dorsal rays. The 25.0 mm. post-larva from Madang has
D. 17 + 20 + 22, (59) and A. 22 4- 12 4- 25, (59). There are 75 lateral
and 14 transverse scale rows and the pigmentation is similar to that
of Australian specimens.

Bregmaceros bathymaster Jordan and Bollman.

Bregmaceros bathymaster Jordan and Bollman 1889, p. 173 (Gulf of
Panama — Type locality).

Bregmaceros longipes Garman 1899, p. 191, pi. 43, figs. 6-9 (Mexico,
Pacific coast near Acapulco). Parr 1931, p. 49.

Bregmaceros macclellandi {now Thompson) Jordan and Evermann 1896-
1900, p. 2526.

D. 18 -f 10 + 19, (44-47). A. 19 + 10 + 19, (44-48). Lat. sc.
60-62. Trans, sc. 10. Depth 6.6 to 7.0, head 5.0 to 5.6 in body length
without caudal. Eye large, 3.0 in head, greater than interorbital and
approximately twice snout. Maxilla extends to or beyond middle of
eye, 2.2 in head. Nuchal appendage 0.3 longer than head. Dorsal fin
inserted slightly in advance of anal fin ; longest ray 0.75 of head length.
Ventrals 0.66 of body length without caudal, extending to end of first
section of anal fin. Pectorals shorter than head. Nape and back brown.
Several rows of dark dots along front part of back and near base of anal
fin. Flanks, cheeks and iris silvery. Dorsal fin dusky. Caudal fin pale,
dusky at base with narrow white cross bar. Other fins pale. (Compiled).
Restricted to the Pacific coast of Central America.

Bregmaceros atlanticus Goode and Bean.

Bregmaceros atlanticus Goode and Bean 1886, p. 165 (West Indies, off
Grenada and Nevis — Type locality ; Gulf of Mexico). Goode and
Bean 1895, p. 389, pi. 95. Jordan and Evermann 1896-1900, p. 2527.
Borodin 1928, p. 13 (Caribbean Sea, off Rancador Reefs). Parr
1931, p. 49. Parr 1937, p. 62 (West Indies, off Cuba and Bahamas).

Bregmaceros macclellandi {non Thompson) ? Norman 1930, p. 339
(Western Africa off Cape Lopez and Sierra Leone). ? Norman 1935,
p. 9 (Angola, off St. Paul de Loanda). ? Fowler 1936, pp. 1254, 1355.

D. (15-16) + x-+ 16, (48). A. (15-16) + (7-8) -f (21-22), (43-50)
or 20 -p 9 + 37, (64). Lat. sc. 65. Trans, sc. 10. Depth 7.6 to 8.0,
head 5.0 to 5.75 in body length without caudal. Eye moderate, 3.5 to
4.0 in head, 1.3 to 1.5 in interorbital, equal to or slightly less than snout.
Maxilla extends to below posterior edge of eye, 2.0 in head. Dorsal fin
inserted directly above anal fin ; longest ray 0.2 greater than head.
Ventrals 0.6 of body length without caudal, extending to end of first
section of anal fin. Pectorals shorter than head. Nuchal appendage
0.5 greater than head, but according to Borodin (1928) twice body length
in young. Body uniformly dusky. Young with many small, dusky
stellate melanophores scattered over body. (Compiled).

An Atlantic Ocean species from Western Indies, Caribbean Sea and
probably the west coast of Africa.

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND

REFERENCES.

Alcock, A. W., 1893. — Natural history notes from H.M. Indian marine survey
steamer “ Investigator.” Ser. II, No. 9. J. Asiatic Soc. Bengal 62 (2) :
169-184, pis. 8-9.

, 1899. — A descriptive catalogue of the Indian deep-sea fishes in the

Indian Museum, collected by the Royal Indian Survey Ship “ Investi-
gator.” Calcutta. 220 pp., 1 map.

Barnard, K. H., 1925. — A monograph of the marine fishes of South Africa. Ann.
S. Afric. Mus. 21 : 1-418, pis. 1-17, 18 figs.

Borodin, N. A., 1928. — Scientific results of the yacht ‘‘Ara ” expedition during
the years 1926-1928, while in command of William K. Vanderbilt. Fishes.
Bull. Vanderbilt Oceanogr. Mus. 1 (1) : 1-37, pis. 3-5, 1 map.

Day, F., 1865. — The Fishes of Malabar. London. 293 pp., 20 pis.

1869. — Remarks on some fishes in the Calcutta Museum. Proc.

Zool. Soc. 1869 : 511-527.

1875-1878. — The Fishes of India. London. Text and Atlas, 778 pp.,

198 pis.

1889. — The Fauna of British India, including Ceylon and Burma

(W. T. Blandford). Fishes, 2. London. 509 pp., 177 figs.

Delsman, H. C., and J. D. F. Hardenberg, 1934. — De Indische Zeevisschen en
Zeevisscherij. Batavia. 388 pp., 273 figs., pis.

Fowler, H. W., 1936. — The marine fishes of West Africa based on the collection
of the American Museum Congo Expedition, 1909-1915. Bull. Amer.
Mus. Nat. Hist. 70 (2) : 607-1493, figs. 276-567.

Garman, S., 1899. — The Fishes. Reports on an exploration off the west coasts

of Mexico, Central and South America, and off the Galapagos Is

“Albatross,” 1891. Mem. Mus. Comp. Zool. Harvard 24 : 1-431, pis. 1-97.

Gilchrist, J. D. F., and W. W. Thompson, 1914. — Descriptions of fishes from the
coast of Natal. IV. Ann. S. A fric. Mus. 13 : 65-95.

1917. — A catalogue of the sea fishes recorded from Natal. II.

Ann. Durban Mus. 1 : 291-431.

Goode, G. B., and T. H. Bean, 1886. — Reports of the results of dredging in

the Gulf of Mexico (1877-78) and in the Caribbean Sea (1879-80), by U.S.
Coast Survey Steamer “ Blake.” Bull. Mus. Comp. Zool. Harvard 12 (5) :
153-170.

— — 1895. — Ocean Ichthyology. Smithson. Contrib. 981 : text and Atlas,

553 pp., 123 pis.

Gunther, A., 1882. — Catalogue of the Fishes in the British Museum. IV. London.
534 pp.

1889.— Report on the Pelagic Fishes. Repts. Sci. Res. “ Challenger,”

31. London. 47 pp., 6 pis.

Jordan, D. S., and C. H. Bollman, 1889. — Descriptions of new species of fishes
collected at the Galapagos Islands and along the coast of the United

States of Colombia, 1887-88 “Albatross.” Proc. U.S. Nat. Mus.

12 : 149-183.

Jordan, D. S., and B. W. Evermann, 1896-1900. — The Fishes of North and Middle
America. Bull. U.S. Nat. Mus. 47 : 1-3313, pis. 1-392.

Jordan, D. S., S. Tanaka, and J. O. Snyder, 1913. — A catalogue of the Fishes
of Japan. Journ. Imp. Coll. Sci. Tokyo 33 (1) : 1-497, figs. 1-396.

Kent, W. Saville-, 1889. — Preliminary observations on a natural history collection
made in connection with the surveying cruise of H.M.S. “ Myrmidon ”

at Port Darwin and Cambridge Gulf in 1888. Proc. R. Soc. Queensl.

6 : 219-242.

McCulloch, A. R., 1926. — Studies in Australian Fishes. No. 8. Rec. Aust. Mus.
15 (1) : 28-39, pi. 1.

Norman, J. R., 1930. — Oceanic Fishes and Flatfishes collected in 1925-1927.
Discovery Repts. 2 : 261-370, pi. 2, figs. 1-47.

1935. — Coast Fishes. Part I. The South Atlantic. Ibid. 12 :

1-58, figs. 1-15.

REVISION OF BREGMACEROS : DESCRIPTIONS OF LARVAL STAGES 53

Okada, Y., 1938. — A Catalogue of Vertebrates of Japan. Tokyo. 412 pp.

Paradice, W. E. T- and G. P. Whitley, 1927. — Northern Territory Fishes. Mem.
Queensl. Mus. 9 (1) : 76-106, pis. 11-15, 3 figs.

Parr, A. E., 1931. — Deepsea fishes from off the Western coast of North and Central
America with keys to the genera Stomias, Diplophos, Melamphaes and
Bregmaceros, and a revision of the Macropterus group of the genus
Lampanyctus. Bull. Bingham Oceanogr. Coll. 2 (4) : 1-53, 18 figs.

1937. — Concluding report on fishes. Ibid. 3 (7) : 1-79, 22 figs.

Richardson, Sir J., 1843. — Ichthyology. The Zoology of the voyage of H.M.S.

“ Sulphur,” under the command of Captain Sir Edward Belcher, during
the years 1836-42 (R. B. Hinds). London : 51-150, 30 pis.

Smith, H. M., 1933. — Contributions to the Ichthyology of Siam. VI. Jouvn.
Nat. Hist. Soc. Siam 9 : 53-87.

Tanaka, S., 1908. — Descriptions of eight new species of fishes from Japan. Annot.
Zool. Japon. 7 (1) : 27-47, 2 figs.

1913. — Figures and descriptions of the fishes of Japan including

Riukiu Islands, Bonin Islands, Formosa, Kurile Islands, Korea and
Southern Sakhalin. II. Tokyo : 187-198, pis. 51-55.

1933. — Fishes. Illustrations of useful, harmful and ornamental

aquatic fauna and flora. Tokyo.

Thompson, W., 1840. — On a new genus of Fishes from India. Charleswovth’s Mag
Nat. Hist. 4 : 184-187, fig.

Tickell, S. R., 1865. — Description of a supposed new genus of the Gadidae, Arakan
( Asthenurus atvipinnis). Jouvn. Asiatic Soc. Bengal 34 (2) : 32, pi. 1.

Weber, M., 1913. — Die Fische der Siboga-Expedition. Leyden. 710 pp., 12 pis.,
123 figs.

Weber M. and L. F. de Beaufort, 1929. — The Fishes of the Indo- Australian
Archipelago. 5. Leiden. ■ 458 pp., 98 figs.

Whitley, G. P., 1941. — Ichthyological notes and illustrations. Austr. Zoologist
10 (1) : 1-50, 32 figs, pis. 1-2.

Wood-Mason, J. and A. W. Alcock, 1891. — Natural history notes from H.M.

Indian Marine Survey Steamer “Investigator.” Ser. II, No. 1. Ann.
Mag. Nat. Hist. (ser. 6) 8 : 16-34.

Vol. LXI., No. 6.

ADDITIONS TO THE FLORA OF
ARNHEM LAND

By C. T. White, Government Botanist, Brisbane.

(Received 28 th October, 1949 ; read before the Royal Society of Queens-
land, 28 th November, 1949 ; issued separately ).

I recently had the pleasure of examining the rich ethno-botanical
collections made in Arnhem Land, Northern Territory of Australia, by
Dr. Donald F. Thomson in 1935-6-7 and in 1941-2-3. The specimens are
preserved in museum jars in the Department of Anthropology at the
University of Melbourne, and in many cases in addition as dried
specimens. In making the determinations I found several species which,
so far as I know, had not previously been collected in Arnhem Land
or other parts of the Northern Territory and two which seem previously
undescribed. A classified account of these new records is offered here-
with. Types of the proposed new species have been deposited in the
Queensland Herbarium.

Family Palmae

Corypha elata Roxb. FI. Ind. ed. 2, 2 ; 176 (1832).

Arnhem Land : Glyde River, D. F. Thomson (photograph only)
June, 1937, only seen growing on watercourses in the valley of the
Glyde River, north-central Arnhem Land (palm 50-60 ft.).

This palm, a native of Bengal and Burma, is widely cultivated
throughout tropical south-east Asia and the Malay Archipelago. It has
been recorded from the , lower Gilbert River, Cape York Peninsula,
Queensland (Beccari ex Ewart and others in Proc. Roy. Soc. Viet,
n.s. 24, pt. II.: 256 (1911) ) but not previously so far as I know from
Arnhem Land. It is probably of Malayan introduction. The deter-
mination is based on a photograph only. It is undoubtedly a Corypha
and I have determined it as above rather than as C. umbraculifera L.,
the Talipot palm, firstly because, as mentioned above, C. elata Roxb.
has already been recorded for Australia and, secondly, because the
photograph shows the spiral furrows on the stem that Blatter (Palms
of British India p. 70) says at once distinguish this species from
C. umbraculifera L.

Family Araceae

Amorphophallus galbra F. M. Bail, in Dept. Agric. Brisbane
Bull. 21 (Bot. Bull. 7) : 68 (1893); Queensl. FI. 5 : 1696, PI. LXXVI
(1902).

North-West Arnhem Land : D. F. Thomson 49, in dry jungle associa-
tions which occur sporadically in suitable pockets near water and among
hills (aroid, approx. 2 ft. high ; fruit orange and bright red in colour ;
very astringent and regarded by natives as poisonous).

Typhonium angustilobum F. Muell. Fragm. Phytogr. Austr.
10: 66 (1876).

North-Central Arnhem Land, near Cape Stewart : D. F. Thomson
39 bis, savannah forest, preferably in fairly damp locations (aroid,
6-12 in. high ; rootstock eaten by the natives).

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND

This was included in the same jar (No. 39) as the more widely spread
and better known T. Brownii Schott, of which, when better known, it
may prove to be only a form or variety.

Family Zingiberaceae

Curcuma australasica Hook. Bot. Mag. t. 5620 (1867).

Arnhem Land : D. F. Thomson 12, jungle associations generally near
water and in sandy soil (flowers during wet season about January, colour
of flower, tinged purplish).

Family Leguminosae
Teptirosia arnhemica sp. nov.

Herba perennis, caulibus paucis humifusis debilibus ca. 0.5 m.
longis simplicibus vel pauciramosis tenuiter pubescentibus. Folia
petiolata, 19-21-foliolata, rachi cum 1-1.5 cm. petiolo 5-8 cm. longa ;
foliola linearia vel lineari-lanceolata, subtus tenuiter pubescentia, apice
apiculata, basi leviter cuneata, breviter petiolulata, nervis praecipuis
ca. 5. Racemi gracillimi, elongati, remotiflori, ad. 18 cm. longi ; flores
pedicellati, pedicellis 2-3 mm. longis, dense strigoso-pubescentibus ;
calyx 2 mm. longus, dense albido-pubescens, lobis acutis ; vexillum extus
dense albido-hirsiitum, unguiculatum, 7 mm. longum et 5 mm. latufn ;
alae glabrae 5 mm. longae et 2 mm. latae ; carina aequilonga ; ovarium
albido-hirsutum. Legumen (immaturum) rectum 3.3 cm. longum, dense
alb: do-hi rsut um .

North Arnhem Land : D. F. Thomson 15, open savannah especially
in sandy soil (herb, 12 in. high ; flowers small, pink or purplish in colour ;
rootstock about the size of a small parsnip, used to poison fish).

Very' close to T. remoti flora F. Muell. ex Benth., but the two can be
distinguished as follows :

Upright shrub or subshrub, leaflets 7-11, oblong-cuneate, lateral veins numerous

and close together T. remotiflora

Herb or subshrub, several weak diffuse stems from a common stock, leaflets 19-21,
linear or linear lanceolate, lateral nerves distant about 5 on each side of the
midrib T. arnhemica

Family Anacardiaceae

Buchanan? a arborescens Blume Mus. Bot. Lugd. Bat. 1 : 183
(1850).

North-East Arnhem Land : D. F. Thomson 113, chiefly in higher
rainfall areas where the vegetation has a rain-forest appearance (tree
15-20 ft.).

Distribution : Burma, Malay Archipelago, Philippine Islands and
tropical Australia.

Buchanania obovata Engler in DC. Monogr. Phan. 4 : 187 (1883).
Far-eastern Arnhem Land : South of Melville Bay and vicinity of

Port Bradshaw ; D. F. Thomson 2, 20 and 81.

Distribution : Confined to Australia.

There has been considerable confusion regarding the. species of
Buchanania in Australia. B. arborescens Bl. as I understand the species
is common in Queensland. According to Dr. Thomson, in Arnhem
Land it grows in the jungle (monsoon forest or light rain-forest) whereas
B. obovata Engl, is a savannah-forest tree. It is rare in Queensland.

ADDITIONS TO THE FLORA OF ARNHEM LAND

Family Sapindaceae

Ganophyllum falcatum Blume Mus. Bot. Lugd. Bat. 1 : 230 (1850).

Arnhem Land : North coast, D. F. Thomson 4, on raised area above
sand beach on fringe of dry jungle (tree 30-35 feet, fruit reddish orange,
matures in December, eaten by natives).

Distribution : Andaman Islands, Philippines, Java, New Guinea
and tropical Australia.

Family Combretaceae
Terminalia carpentariae sp. nov.

Arbor 10-13 m. alta, ramulis densissime velutino-pubescentibus-
Folia subchartacea oblonga vel rarius elliptico-oblonga plerumque
obtusissima et interdum leviter emarginata, rarissime breviter acuminata,
basi obtusa vel rarissime brevissime cuneata, utrinque dense et molliter
pubescentia, nervis praecipuis ca. 7 in utroque latere, reticulatione
utrinque prominulo vel subtus interdum plus vel minus prominenti ;
petiolus 2-4 cm. longus ; lamina 8-12 cm. longa, 6-9 cm. lata. Spicae
fructiferae 6-8 cm. longae, densissime velutino-pubescentes. Drupae
dense velutino-tomentosae, 3 cm. longa6, 1.7 cm. latae, 1 cm. diam.,
ellipsoideae, rostratae compressae vel plano-convexae, lateribus acute
angulatis.

Northern Territory : Arnhem Land : north coast, Crocodile
Islands, D. F. Thomson 111 (type), chiefly in zone fringing the sea-front
(tree 30-40 feet, cambium layer used for caulking canoes). Settlement
Creek, L. J. Brass 236, October, 1922, hill country (small tree, fruit
said to be excellent eating when stewed. Local name “ Plum Tree ”).

Queensland : Burke District : Gulf of Carpentaria, Mornington
Island : J. F. Bailey, June 1901 ; E. W. Bick 236, October, 1922 ; Lawn
Hill : H. I. Jensen 94, May, 1940.

This tree grows in several parts of the “ Gulf ” country of Queens-
land and is apparently common and widely spread in the Northern
Territory, as in addition to the specimens quoted above it is represented
by several sheets in the Blake (Northern Australia Regional Survey) and
Specht (Australian-American Arnhem Land Expedition) collections. It
is undoubtedly very closely allied to T. platyphylla F. Muell., with which
it has been confused in the past. Another very closely allied species
is T. aridicola Domin.

The Australian species of Terminalia are notoriously difficult to
delimit, but I think we are dealing with three distinct species here which
can be keyed out or rather summarised as follows :

Leaves mostly cuneate, rarely subobtuse at the base, more or less densely
pubescent on both surfaces, petiole 1-2 cm. long, lamina 4-8 cm. long,
2-5 cm. wide, lateral nerves about 6 on each side of the midrib ; drupe
broadly and shortly ellipsoid, not rostrate, not compressed but with
sharp angles almost developed into lateral wings in the younger stage, dis-
appearing and only remaining as a sharp edge in the mature fruit, densely
pubescent, 2.5 x 2 x 1.5 cm. T. aridicola

Leaves mostly cuneate, very rarely obtuse at the base, glabrescent above or at
most thinly pubescent, petiole 2-3 cm. long, lamina 10-17 cm. long, 6-10 cm.
wide, lateral nerves 8-10 on each side of the midrib ; drupe rostrate, narrowly
ellipsoid without any angles or wings, not compressed nor inclined to be plano-
convex, thinly pubescent, 3xlxl cm T. platyphylla

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND

Leaves mostly obtuse, , very rarely indistinctly and very shortly cuneate at the
base, densely velvety pubescent on both faces, petiole 2-4 cm. long, lamina
8-12 cm. long, 6-9 cm. wide, lateral nerves 7 on each side of the midrib ; drupes
rostrate, ellipsoid, more or less compressed or plano-convex (or at least very
slightly convex on one face and markedly so on the other), acutely angled on
the sides, densely tomentose, 3 x 1.7 x 1 cm T. carpentariae

Family Thymelaeaceae

Phaleria blumei Benth. var. latifolia Benth. FI. Austr. 6 : 38
(1873).

Arnhem Land : Caledon Bay, D. F. Thomson 57, August 1936, near
the beach (shrub, used as a fibre plant).

Distribution : Malay Archipelago, tropical Australia.

Family Rhizophoraceae

Bruguiera parviflora (Roxb.) Wight & Arn. Prodr. 311 (1834).

North Arnhem Land : Crocodile Islands, D. F. Thomson 28, Septem-
ber 1935, mangrove zone (tree 20-25 feet, wood used by natives for
canoe paddles).

Distribution : India, Malay Archipelago, tropical Australia.

Vol. LXI., No. 7.

HEAVY MINERAL BEACH SANDS OF

SOUTHERN QUEENSLAND.— Part II.

PHYSICAL AND MINERALOGICAL COMPOSITION,
MINERAL DESCRIPTIONS, AND ORIGIN OF THE
HEAVY MINERALS.

By A. W. Beasley, M.Sc., Ph.D., D.I.C., F.G.S., c /- Department of
Geology, University of Queensland.

(With Five Text-Figures and Six Plates.)

(Received ZQth August, 1949 ; tabled before the Royal Society of Queens-

land 28 th

November, 1949 ; issued separately — - — - —

- )•

CONTENTS.

Page

Summary

Introduction

II.

Places of Collection of the Beach Sand Samples

III.

Mechanical Composition of the Natural and Panned

Heavy

Mineral Beach Sand Concentrates

IV.

Mineralogical Methods ... ... ...

Mineralogical Composition of the Heavy Mineral Beach Sand

Concentrates

VI.

Description of the Minerals

VII.

Geographic Distribution of the Heavy Minerals

VIII.

Origin of the Heavy Minerals

IX.

Conclusions and Outline of Geological History of the Heavy

Minerals ... ...

101

Acknowledgments ...

103

XI.

Bibliography

104

SUMMARY.

The physical and mineralogical composition of 50 samples of heavy
mineral sands, collected from along a 300-mile stretch of the Eastern
Australian coast, are described. Results of sieve analyses of the samples
of natural concentrate, and of the panned heavy mineral concentrate
obtained from them, are presented. The median diameter, coefficients
of sorting and log skewness of the heavy mineral concentrates are given,
and the values plotted against distance of the samples along the coast
from south to north. Descriptions and comparisons of physical com-
positions are based mainly on these measures, and they are shown to be
of use in suggesting places of heavy mineral addition and direction of
transport along the coast. The percentage of heavy minerals in the
natural concentrates ranges up to 95.2% by weight. Mineral analyses
of the heavy mineral concentrates are given in weight percentages, and
20 species are listed. The minerals are described and the geographic
distribution along the coast is discussed. Over 90% of the heavy minerals
in all the samples consist of zircon, rutile and ilmenite. Decreases in
the degree of zircon abrasion around the major coastline breaks and
about the headlands of Mesozoic sandstone suggest that heavy mineral
material has been added to the shore at these points. The geology of
the region, and study of the heavy mineral assemblages of 20 selected
rock samples and 1 1 river sand samples indicate that Mesozoic freshwater
sandstones are the immediate source rock for most of the heavy minerals
in the beach sands. The main primary sources are Permian granitic

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND

rocks, but some of the ilmenite in the sands has been derived from
Tertiary basalts. An outline of the geological history of the heavy
minerals is presented.

I. INTRODUCTION.

From 1945 to 1947 an investigation was carried out into the nature,
distribution, extent, and manner of formation of the heavy mineral
sand deposits of the S.E. Queensland coast. The results of this have
already appeared in these Proceedings (Beasley, 1948).

During the field work samples were collected from places of heavy
mineral concentration on the beaches and the adjacent dunes. These,
together with a number collected by the Queensland Geological Survey,
form the material on which the present work is based. The samples
were obtained from the 250-mile stretch of Queensland coastline between
the New South Wales border in the south and Indian Head on Fraser
Island in the north. A small representative collection was obtained from
Northern New South Wales for comparison with the Queensland sands.
These were collected along the coast from the State border southwards
for 50 miles to Ballina.

The black sand deposits of commercial importance in Eastern
Australia occur between Ballina and Moreton Island. From Moreton
Island northwards to Fraser Island the deposits are small and widely
separated.

Almost all sand samples were obtained by boring with a 4-inch
post-hole digger. Samples of black sand seams and composite samples
of entire bores were taken as previously described (Beasley, 1948, p. 118).
A small number of samples was obtained from cased bores put down
with hand and power-driven plants, and a few were obtained from the
faces exposed in the workings of operating companies.

The main objects of the present work have been to determine the
physical and mineralogical constitution of the sands, particularly the
heavy mineral content, to describe the heavy minerals, and to enquire
into their origin. For economic reasons, weight percentages of the
heavy mineral species have been determined in preference to mineral
grain number percentages.

II. PLACES OF COLLECTION OF THE BEACH SAND SAMPLES.

The localities given are numbered consecutively from south to north.
Unless otherwise stated, the samples are from black sand seams in bores.
The first six samples are from Northern New South Wales.

1. Top of beach just S. of mouth of Richmond River, Ballina.

2. Immediately in front of foredune on S. side of Lennox Head.

3. Seam exposed in workings on Seven-Mile Beach, just S. of Byron Bay.

4. Seam exposed in beach workings near mouth of Crabbe's Creek, 4 miles N. of
New Brighton.

5. Behind foredune, £ mile S. of Norries Head.

6. Top of beach, 3 miles S. of Cudgen Headland.

7. Top of beach 4 chains N. of Tugun Surf Pavilion.

8. Seam exposed in beach workings at Flat Rock, Tugun.

9. Seam exposed in beach workings £ mile S. of South Nobby Headland and
opposite Fifth Avenue, Burleigh.

10. Composite sample of 6 feet in bore in most landward of Recent coastal dunes,
\ mile inland, at South Nobby.

11. Immediately behind third dune ridge inland from beach, £ mile N. of North
Nobby.

TEXT- FIGURE I.— LOCALITY MAP, SHOWING PLACES OF

MINERAL BEACH SANDS OF SOUTHERN QUEENSLAND, Part II. 61

12. Hollow between two dupe ridges £ mile inland, 1 mile S. of Broadbeach Surf
Pavilion.

13. Seam exposed in workings immediately behind foredune, £ mile S. of Broad-
beach Surf Pavilion.

14. Immediately in front of foredune, on Southport sandspit, 2 miles S. of Nerang
River mouth.

15. Seam exposed in vertical wave-eroded scarp at top of beach, £ mile N. of
southern extremity of South Stradbroke Island.

16. Composite sample of 3 ft. bore containing thin black sand seams, immediately
in front of foredune, 3^ miles N. of southern extremity of South Stradbroke
Island.

17. Top of beach, 5 miles N. of southern extremity of South Stradbroke Island.

18. Seam exposed in deep hollow immediately in front of foredune, miles N. of
southern extremity of South Stradbroke Island.

19. Top of beach near extreme southern end of North Stradbroke Island, just N.
of Jumpinpin Break.

20. Foot of foredune, 15 miles S. of Pt. Lookout, North Stradbroke Island.

21. Foot of foredune, 12 miles S. of Pt. Lookout. North Stradbroke Island.

22. Seam exposed in deep hollow or “ blow-out ” in foredune, 9£ miles S. of Pt.
Lookout, North Stradbroke Island.

23. Composite sample of 10 ft. bore sunk through wind-concentrated heavy mineral
sand, just seaward of crest of Pleistocene foredune, i mile S. of mouth of Blue
Lake Creek, North Stradbroke. Island.

24. Composite sample of 18 ft. bore sunk through wind -concentrated heavy mineral
sand, in high Pleistocene dunes 1 mile inland from present strandline and
\ mile S. of Blue Lake Creek, North Stradbroke Island.

25. Composite sample of 18 ft. bore sunk through wind-concentrated heavy mineral
sand near Blue Lake, in region of Pleistocene dunes, miles inland from
present strandline.

26. Composite sample of 36 ft. bore in Eighteen Mile Swamp adjacent to western
margin, \ mile N. of Blue Lake Creek, North Stradbroke Island.

27. Composite sample of 18 ft. bore sunk through wind-concentrated heavy mineral
sand, at elevation of 280 feet, near crest of Pleistocene foredune, 1^ miles N.
of Blue Lake Creek, North Stradbroke Island.

28. Top of beach, £ mile S. of Pt. Lookout, North Stradbroke Island.

29. Top of beach, 1 mile E. of Rocky Point, on northern side of Stradbroke Island,
just W. of Pt. Lookout.

30. Surface accumulation of black sand on beach at Amity Point Wharf, Strad-
broke Island.

31. Top of beach, miles N. of southern end of Moreton Island.

32. Top of beach, 10 miles N. of southern end of Moreton Island.

33. Top of beach, 14 miles N. of southern end of Moreton Island.

34. Top of beach, 4£ miles S. of Cape Moreton, Moretop Island.

35. Top of beach, 1£ miles S. of Cape Moreton, Moreton Island.

36. Top of beach, \\ miles W. of North Pt., near Yellow Patch, Moreton Island.

37. Top of beach, 1 mile E. of Comboyuro Pt., Moreton Island.

38. Immediately in front of foredune, 3 miles S. of northern end of Bribie Island.

39. Black sand surface accumulation on beach adjacent to Caloundra Head.

40. Black sand surface accumulation at top of beach near Alexandra Headland.

41. Immediately in front of foredune, J mile S. of Pt. Arkwright.

42. Immediately in front of foredune, 1 mile S. of Paradise Caves, Noosa, along
Coolum Beach.

43. Black sand surface accumulation on beach, 10 chains S. of mouth of Noosa
River.

44. In front of foredune, 3 miles N. of mouth of Noosa River, along Laguna Beach

45. On Laguna Beach, 10 miles N. of Noosa.

46. Immediately in front of foredune, £- mile S. of Double Island Point.

47. Black sand surface accumulation on beach, 1£ miles S. of Inskip Point.

48. Black sand surface accumulation on beach, 1 mile N. of Hook Point, Fraser
Island.

49. Black sand surface accumulation on beach, immediately S. of Poyungan Rocks
(recently cemented beach sand).

50. Immediately in front of foredune, 1 mile S. of Indian Head, Fraser Island.

The localities are shown in Text-figure 1.

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND

III. MECHANICAL COMPOSITION OF THE NATURAL AND
PANNED HEAVY MINERAL BEACH SAND CONCENTRATES.

Mechanical analysis of the samples of natural concentrate was carried
out to obtain knowledge of their size distribution. As the grains generally
have diameters greater than 0.062 mm. (1/16 mm.), it was possible to
separate them into fractions with sieves.

Sieve analysis of samples of the heavy mineral concentrate obtained
from the natural concentrate by panning also was carried out. Know-
ledge of the size distribution, sorting, and skewness of these samples
was considered desirable for purposes of comparison. Satisfactory
comparisons could not be made from the mechanical analysis of the
natural concentrate owing to the marked difference in specific gravity
between the light and heavy minerals and the local variations in the
degree of natural, heavy mineral concentration. However, as in all the
samples of panned concentrate over 90% of the minerals was found to
consist of zircon, rutile and ilmenite, which have similar specific gravities
and have been naturally concentrated in grains of very similar size,
it has been possible to compare and describe these concentrates using
statistical measures derived from the cumulative frequency , curves.
An interpretation of the mechanical analyses in terms of heavy mineral
supply and transportation along the coast has tlius been possible.

From Ballina to Indian Head the ocean coastline consists of a
series of arc-shaped sandy beaches separated by rocky headlands. These
coastline curves are broken in places by river mouths and breaks between
the coastal islands, and between the islands and the mainland. The
beaches are gently sloping, ranging in width up to 200 feet at low tide,
and are bordered by a belt of coastal dunes. A detailed account of
the physiography has been given elsewhere (Beasley, 1948, pp. 111-116).
In the southern half of the area the headlands are chiefly composed of
Lower Palaeozoic slates and greywackes and Tertiary basalts ; but from
Cape Moreton northwards they are largely of Mesozoic sandstones, except
for Indian Head, which is composed of Tertiary basalt. Throughout
and beyond the area, the prevalent wind is the South-East Trade, and
close to the land there is an inshore ocean current setting in a northerly
direction with a rate of from a quarter to one knot. In the summer months
south-east gales are not infrequent and, with the powerful waves striking
the beach obliquely, the sand is drifted along the beach. Under these
influences, combined with the longshore ocean current, it would seem
that the direction of sand transport is mostly northward.

Mechanical Analysis.

In the laboratory, the samples of natural concentrate, usually of
the order of several hundred grams, were washed free of salt, dried, and
split by a rotary sample-splitter to approximately 40 grams. The sample-
splitter consisted of a turntable with a tin mounted on it containing a
number of glass tubes, into which the sand was discharged from an
overhead funnel. The split samples were weighed, then shaken in a
nest of sieves with a Ro-tap mechanical shaker for 30 minutes. The
sieves used were numbers 60, 85, 100, 120, 150 and 200 of the British
Standard Series, the apertures respectively being 0.251, 0.178, 0.152,
0.124, 0.104 and 0.076 mm. Distortion of the mesh due to wear was
negligible. The resulting size fractions were then weighed, percentages
calculated, and the results tabulated (Table I).

(The abbreviations “ Nat.” and “ Pan.” immediately after the sample numbers in the following Table refer respectively to the natural
concentrate and the panned concentrate derived from it. In a few cases the panned concentrate only was sieved.)

MINERAL BEACH SANDS OF SOUTHERN QUEENSLAND, PART II.

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MINERAL BEACH SANDS OF SOUTHERN QUEENSLAND, PART II.

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66 PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND

To obtain heavy mineral samples from the natural concentrates for
sieve analysis, panning was found to be satisfactory as the heavy minerals
are abundant and over 95% of them differ markedly in specific gravity
from the light minerals. The samples of natural concentrate were
weighed and panned ; porcelain evaporating dishes were used in the
final stages instead of prospecting dishes as the heavy minerals remaining
showed up more plainly against the white background. The panned

Table II. — First and Third Quartiles, Medians, Coefficients of Sorting
and Log Skewness of the Panned Concentrates.

No.

Millimetres

Log Sk

1 . .

.120

.115

.108

1.05

.004

.120

.115

.108

1.05

.004

3 . .

.118

.112

.104

1.06

.004

.120

.115

.108

1.05

.004

5 . .

.120

.113

.106

1.06

.000

.122

.114

.106

1.07

.000

7 . .

.122

.114

.106

1.07

.000

8 . .

.122

.114

.107

1.07

-.001

.120

.113

.106

1.06

.000

10 . .

.190

.149

.112

1.30

.000

11 . .

.124

.115

.107

1.08

-.001

12 . .

.120

.111

.103

1.08

-.001

13 . .

.122

.114

.107

1.07

-.001

.122

.113

.105

1.08

-.001

.192

.124

111

1.31

-.080

.136

.115

.105 ’

1.14

-.017

17 ..

.120

.110

.100

1.10

.000

.149

.118

.110

1.16

-.036

.160

.120

.111

1.20

-.048

.147

.133

.118

1.11

.004

.155

.114

.108

1.20

-.055

.120

.111

.100

1.10

.004

.120

.110

TOO

1.10

.000

.120

.110

TOO

1.10

.000

.121

.110

TOO

1.10

-.001

.123

.112

.104

1.09

-.005

.120

.110

TOO

1.10

.000

.120

.111

TOO

1.10

.004

.154

.117

.107

1.20

-.042

.160

.116

.106

1.23

-.052

31 ..

.180

.151

.119

1.23

.009

.123

.115

. .108

1.07

-.001

.148

.116

.106

1.18

-.034

.123

.113

.104

1.09

-.001

.166

.123

.111

1.22

-.046

.164

.122

.111

1.21

-.046

.153

.118

.110

1.18

-.043

.190

.153

.120

1.26

.005

.230

.200

.171

1.16

.000

.139

.114

TOO

1.18

-.034

.217

.180

.151

1.20

-.008

.172

.150

.118

1.21

.019

.164

.128

.113

1.20

-.030

.121

.114

.106

1.07

.004

.124

.115

.106

1.08

.000

.150

.118

.110

1.17

-.038

.173

.140

.114

1.23

— .006

.151

.118

.109

1.17

-.037

.120

.111

TOO

1.10

.004

50 . .

.131

.117

.110

1.09

-.011

MINERAL BEACH SANDS OF SOUTHERN QUEENSLAND, PART II. 67

heavy mineral concentrate was then dried and weighed, and the weight
percentage of heavy minerals in the sample of natural concentrate was
determined (see Table I). Each sample of panned concentrate was split
by the rotary sample-splitter to approximately 40 grams, and the split
samples were weighed and screened as described for the natural
concentrates. The resulting size fractions were weighed, percentages
calculated, and the results tabulated (Table I). Cumulative frequency
curves were constructed from this information, and from them the first
,and third quartiles and the median diameter were tabulated (Table II).

Comparison of the samples is based on the median diameter and
the coefficients of sorting and log skewness, following Trask (1932).
Where Q1 and Q3 are the first and third quartiles, respectively, and M
the median, the coefficient of sorting, is -y/Ql/QS. It expresses the
measure of the average quartile spread. Thus perfect sorting equals
unity, and the larger the value the more poorly sorted is the sample.
The coefficient of skewness, a measure of the dis-symmetry of the size
distribution with respect to the median, is derived from the expression
log Q1 x log Q3/(log M)2.

For convenience the logarithm of the skewness is used. Thus a minus
value indicates that the mode or peak of the simple frequency curve
is on the coarse side of the median, while a positive value indicates the
opposite. The coefficients of sorting and log skewness of the panned
concentrates were calculated and tabulated (Table II).

Graphical Representation and Discussion of Results.

From an examination of the mechanical analyses of the natural
and panned heavy mineral concentrates shown in Table I it will be
seen that the light constituents (essentially quartz) occur in coarser
grains than do the heavy minerals. This may be explained by the fact
that for a certain size of quartz there is a smaller size of heavy mineral
which is deposited with it, because they have the same “ hydraulic
value ” or the same settling rates. Generally, the greater the weight
percentage of heavy minerals in the natural concentrate, the less the
amount of material retained on the two coarsest sieves, B.S.S. Nos. 60
and 85. In all except three of the samples of panned heavy mineral
concentrates, the maximum sieve-fraction percentage, which ranges from
32% in No. 31 to 72% in No. 4, lies in the 0.124 to 0.104 mm. grade size.
The three exceptions are samples No. 38, 39 and 41, all of which have
the maximum sieve-fraction percentage, ranging from 30.4% in No. 38
to 6,3% in No. 39 in the 0.251 to 0.178 mm. grade size.

In Text-figures 2, 3 and 4 the median diameter, coefficients of
sorting and log skewness, respectively, of the samples of panned con-
centrate have been plotted against distance along the coast, from south
to north.

Median Diameter. — Table II and Text-figure 2 show that the median
diameter of the panned heavy mineral concentrates ranges from a
minimum of' 0.110 mm. (Stradbroke Island) to a maximum of 0.200
mm. (Caloundra).

From Ballina northwards to the South Passage the trend is for a
slight decrease in the median values, apart from abrupt increases around
the two major coastline breaks in this stretch (Nerang River mouth
and Jumpinpin Break). These increases are much more pronounced

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND

on the northern side of the breaks, but on neither side are they main-
tained for any great distance. The general trend for a slight decrease
in median values northwards in this region suggests that the material
has come mainly from the south. The decrease is in the direction of
the beach drift. The sudden, temporary increases around the Nerang

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River mouth and the Jumpinpin Break suggest that heavy mineral
material is added at these points. The fact that there is much less
increase on the southern side of these two major breaks appears to be
due to the northward direction of sand transport. In this 106 mile-long
stretch of coast the median diameter of the panned heavy mineral
concentrates ranges from a minimum of 0.110 mm. to a maximum of
0.133 mm.

MINERAL BEACH SANDS OF SOUTHERN QUEENSLAND, PART II. 69

It is of- interest to find that the median diameter of one of the five
dune sand concentrates, sample No. 10, is as large as 0.149 mm. This
may be because the heavy minerals of this sample from a low, inland
dune have not been through the surf zone as often, and have not been
subjected to as much abrasion by water, as those in the more recent
beach sand deposits adjacent to the present strandline. Some 40 miles
further north, the median diameter of the other inland dune sand
concentrates, samples 23, 24, 25 and 27 is only 0.110 mm. These four
concentrates, however, are from the phenomenally high dunes of North
Stradbroke Island, and were collected from bores sunk at elevations
ranging from 50 to 280 feet above present sea-level, and from as far as
1J miles inland from the present strandline. The much smaller median
diameter than that of sample 10 apparently is related to the stronger
winds which formed these unusually high dunes. Greater distance from
the source of the heavy minerals may also be a contributing factor.
Alf five of these concentrates are from fixed dunes in which there has
been no sand-movement for many years. They have been included in
this study for comparison with the 45 beach sand samples, as they are
from low-grade dune sands which are at present attracting some
commercial interest.

From the South Passage northwards to Indian Head the median
diameter shows marked increases in samples from most of the headlands,
as well as just north of two major coastline breaks (South Passage and
North Passage). In this region, the headlands of Cape Moreton,
Caloundra Head, Point Arkwright, Noosa Head and Double Island Point
are made up largely of Mesozoic sandstones* while Indian Head is com-
posed of Tertiary basalt. As the black sand deposits from the South
Passage northwards are of small extent and are usually restricted to the
vicinity of the headlands, it seems that the source of these heavy minerals
with large median diameter is comparatively local in most cases. No
general trend in median values is apparent for this 190 mile-long stretch
of coast. From Caloundra Head northwards, however, the amount of
increase in median diameter at succeeding headlands generally diminishes.

Sorting. — The heavy mineral concentrates are well sorted, the
coefficient of sorting ranging from 1.05 to 1.31 (Table II and Text-
figure 3). The least well-sorted heavy minerals usually occur just north
of major coastline breaks and about the headlands of Mesozoic sandstone.
It will be noticed too that the degree of sorting increases markedly away
from these coastline breaks and sandstone headlands.

From Ballina to the mouth of the Nerang River, an area in which
there are no headlands of Mesozoic sandstone and no major coastline
breaks, there is a slight decrease in the degree of sorting (that is, an
increase in the coefficients of sorting). This is unexpected, for throughout
the area the direction of sand drift is northward under the influence of
the prevailing SE. winds and northerly longshore current. Although
the decrease is only a very small one, sorting was expected to increase
in the direction of drift.

With reference to the five dune sand concentrates studied, sample
number 10, which has a comparatively large heavy mineral median, is
the least well-sorted, while samples 23, 24, 25 and 27 (those from North
Stradbroke Island), which have small median diameters, are all very
well sorted. However, the dune sand concentrates do not have the
highest degree of sorting of the samples studied. Text-figures 2 and 3

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND

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O W

u- L

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MINERAL BEACH SANDS OF SOUTHERN QUEENSLAND, PART II.

TEXT- FIGURE 4.— LOG SKEWNESS OF PANNED CONCENTRATES
PLOTTED AGAINST DISTANCE ALONG COAST.

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND

show that poorer sorting of the individual samples is often paralleled
by a large median, and greater sorting by a smaller median. That is,
the sorting generally improves with increase in fineness (decrease in
median diameter).

Skewness. — The values for skewness (Table II and Text-figure 4)
range from — .080 (just north of the Nerang River mouth) to -f .019
(at the Noosa end of Coolum Beach). The sample with the highest
negative skewness is the least well-sorted of all the samples studied.
Text-figure 4 indicates the very small values for skewness in the region
from Ballina to the mouth of the Nerang River. Thus there are nearly
symmetrical simple frequency curves for the concentrates from this
stretch of coastline, with the mode very close to the median and actually
corresponding with it in five samples.

From the Nerang River mouth northwards to Indian Head the
mode usually is on the coarse side of the median. In this area there are
only eight samples with positive values for log skewness. Four of the
five dune sand concentrates examined show no skew, while the remaining
one (sample 25) has a very small negative skewness of —.001. Apart
from the above, no general trends are apparent in the skewness values.
The fact that the mode usually is on the coarse side of the median north-
wards from the Nerang River mouth again suggests that the heavy
minerals have been contributed more recently to the beach than those
in the southern part of the region.

IV. MINERALOGICAL METHODS.

The methods adopted in determining the mineralogical composition
of the sand and rock samples studied in this investigation are described
below, as well as the methods of determining abrasion and grain size
measurement under the microscope. River-sand and rock samples were
examined in connection with the enquiry into the origin of the beach
sand heavy minerals. In all cases the rotary separator already described
was used for sample splitting, and bromoform of specific gravity 2.86 was
employed for the initial heavy liquid separation.

Sand Samples.

A small sample of the natural concentrate, split from the bulk
sample, was submitted to bromoform separation in a nearly cylindrical
funnel to. minimize adherence of particles to the sides. After washing
with industrial methylated spirits and drying in an oven, the heavy
mineral concentrate was split in size to approximately 5 grams and
weighed. As it was desired to distinguish between magnetite, ilmenite,
chromite, and “ black ” rutile grains, the highly magnetic and moderately
magnetic minerals were separated from the samples with an electro-
magnet. A number of black and nearly black rutile grains was found
in all the samples, identification being established definitely by chemical
analysis of hand-picked grains. The highly magnetic fraction (magnetite)
was extracted first with the electromagnet calibrated for the purpose,
and its weight percentage determined. An aluminium shield was placed
under the pole-pieces to make removal of the magnetite grains easier
after the current was switched off. The moderately magnetic minerals
(ilmenite, chromite and garnet) next were removed with the electromagnet
precisely calibrated, and the weight percentage of this fraction was
determined. The garnet (almandine) was separated from the ilmenite

MINERAL BEACH SANDS OF SOUTHERN QUEENSLAND, PART II.

and chromite with concentrated Clerici’s Solution of specific gravity
4.25 and, after washing with water and drying, the weight percentages
of both the garnet and the ilmenite-chromite fractions were determined.
In the case of the beach sand samples, the small amount of chromite
present was separated from the ilmenite with concentrated Clerici’s
Solution heated to 42° C. (at which temperature its specific gravity is
about midway between that of the chromite and the ilmenite), and the
percentage of each of these minerals in the sample was calculated. In
the river sand samples, however, the small amount of chromite, where
present, was not separated from the ilmenite.

The quantity of weakly magnetic minerals was very small and they
were not separated from the non-magnetic minerals in the sample. The
combined weakly magnetic and non-magnetic fraction of the sample
was weighed, and then divided into two size fractions by shaking in a
B.S.S. No. 120 sieve (aperture size 0.124 mm.) until particles ceased to
pass through the mesh. The weight percentages of the resulting size
fractions were calculated, and each fraction was then split to about
one to two thousand grains and mounted entirely on a glass slide. In
most cases permanent mounts were made in Canada balsam, Twenhofel
and Tyler’s (1941, p. 168) technique being followed. Temporary mounts
in eugenol (clove oil) were also made. The grains of each mineral species
were then counted in some 8 to 12 different fields across different parts
of the mount, the number of grains usually being of the order of 800
to 1,000. The number of grains of each mineral species was multiplied
by the specific gravity of the mineral in order to obtain a figure in terms
of weight. The specific gravity of the zircon, rutile, monazite and
cassiterite was determined with a pycnometer as 4.68, 4.21, 5.19 and 6.90
respectively. For these determinations, small quantities of the com-
mercial mineral concentrate, after handpicking under a binocular micro-
scope to obtain purity, were used. The specific gravity of the tourmaline-
leucoxene, epidote, spinel, corundum, hypersthene, andalusite, horn,
blende, sphene, staurolite, and kyanite was taken as 3.1, 4.0, 3.4, 3.6,
4.0, 3.4, 3.1, 3.2, 3.5, 3.7, and 3.6 respectively. The percentages of the
various mineral species in each of the size fractions were then calculated.
These figures were each multiplied' by the weight percentage of the size
fraction divided by 100, and the results for like species in both size
fractions were added. The percentages of these various minerals in the
entire sample were then determined from multiplication by the weight
percentage of the weakly and non-magnetic mineral fraction divided
by 100. Because of the small spread of all the weakly-magnetic and
non-magnetic minerals present, it was possible to take the grain size
variation into account by division into two fractions with the B.S.S.
No. 120 sieve in the determination of the weight percentages of these
minerals.

Rock Samples.

In most cases the size of the samples of metamorphic, igneous and
sedimentary rocks taken for breaking down was about one-quarter the
size of a normal rock hand-specimen.

Metamorphic and Igneous Rocks. — The samples of metamorphic and
igneous rocks were mechanically disintegrated, first by cracking in a
jaw-cracker, and then by hammering in an iron mortar with closely
fitting pestle, similar to that figured by Krumbein and Pettijohn (1938,
p. 313). Disintegration was continued until the particles obtained were

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND

monomineralic. To avoid the formation of a large amount of fine rock-
flour and total destruction of the original form of the grains, the material
was sieved at intervals, the oversize being crushed until it was reduced
to grains consisting of single minerals. With the plutonic rocks the
sieve used to separate the composite from the monomineralic particles
was B.S.S. No. 52 (aperture size 0.295 mm.), while B.S.S. No. 85 (aperture
size 0.178 mm.) was used with the met amorphic and volcanic rocks.
After the removal of the rock-flour by washing with distilled water and
decanting, the material was dried, split to about 50 grams, and weighed.
The heavy mineral particles were separated in bromoform and, after
weighing, the index-figure was calculated. This is the weight percentage
of mineral grains of specific gravity greater than 2.86 obtainable from
the crushed rock material which has been washed free from rock-flour.
The magnetic minerals were then separated with the electromagnet.
Because of the variety and large bulk of magnetic minerals in the igneous
rocks and the difficulty of effecting clean separations, the weight per-
centages of highly magnetic, moderately magnetic, and weakly magnetic
mineral fractions were not determined individually. In most cases,
however, a rough separation into these three magnetic groups was made
as it facilitated the identification of certain of the minerals and assisted
in estimating the relative abundance of the minerals in the samples.
The magnetic and non-magnetic heavy mineral fractions were examined
under the microscope in permanent and temporary mounts. Because
of the disintegration of the rocks entirely by mechanical means, it was
considered impracticable to determine the mineral percentages by any
method employing grain counting. The relative abundance of the
minerals, accordingly, were determined by estimation and recorded by
symbols in the Milner (1929, p. 386) Scale.

Sandstones. — The sandstone samples were broken into small pieces
in the jaw-cracker, and then heated to a high temperature. While hot
they were dropped into a beaker of cold water, and allowed to remain
in it for several days. In some cases the material thus became partly
broken down, and the individual particles were freed by gentle crushing
with a pestle or with the fingers. In other cases the rock was disintegrated
by warming with a 20% solution of HC1. To restrict the time of acid
treatment, the material was disturbed and partly broken down with a
pestle at intervals during the digestion. Vigorous crushing was avoided,
however, so that the grains would be as nearly as possible in the same
condition as before disintegration. After the sample was completely
disintegrated, the mineral particles were washed and weighed, the heavy
minerals separated in bromoform, and the weight percentages determined.
These correspond to the index-figure of the other rocks. Because of the
small bulk of the heavy minerals obtained from each sample, it was
impracticable to divide it into fractions with the electromagnet. In
some cases the bulk was such that the heavy minerals were mounted
entirely on a glass slide. In other cases, where the bulk was greater,
part of the fraction was' mounted on a glass slide and the remainder
was kept unmounted for supplementary study. The grains of each
mineral species were then counted in the same way as that described
above for the weakly magnetic and non-magnetic fraction of the sand
samples, and the weight percentages of the various minerals were deter-
mined. Unfortunately, it was impossible to distinguish with certainty
between the black iron ores under the microscope. However, examination
of the unmounted material showed the iron ores to be moderately

MINERAL BEACH SANDS OF SOUTHERN QUEENSLAND, PART II.

magnetic. From this fact, combined with an examination of these
grains by reflected light, it would appear that they are almost entirely
ilmenite, although very small amounts of chromite and magnetite are
sometimes present. In this work the iron ore grains were all multiplied
by the specific gravity of the ilmenite concentrate, determined with a
pycnometer as 4.69, and the specific gravity of each of the various other
minerals was the same as that used for the sand samples. Because of
the small bulk of heavy minerals, allowance was not made for the varia-
tion in size of the mineral grains in the determination of the weight
percentages. However, it was observed under the microscope that the
variation in size of the heavy mineral grains was not great. Accordingly,
it is felt that a fairly high degree of accuracy can be assigned to the
mineral weight percentages calculated from the number of grains of
each mineral species and its specific gravity.

Abrasion.

For each of the sand samples and the metamorphic and sedimentary
rocks a quantitative determination of abrasion was carried out by calculat-
ing the number percentage of rounded zircon grains as distinct from
euhedral and subhedral grains. The total number of zircon grains
examined and counted in each, sample was of the order of 300.
Euhedral grains are those with all visible crystal faces and edges intact,
while subhedral grains are those with only some faces and edges
recognizable, and rounded grains those with no faces or edges
identifiable. Zircon was chosen as a standard for this quantitative
abrasion work because of its great stability, its abundance in the
samples, and the wide range in the degree of its abrasion due to a hardness
greater than most of the other heavy minerals present.

Grain Size Measurement.

As the bulk of heavy minerals obtained from the rock samples was
insufficient for sieve analysis, measurements were made with an eyepiece
micrometer of the zircon grains in each of these assemblages. For this
work the intermediate diameter of 100 zircon grains in each sample was
determined and a mean taken in each c,ase, following Allen (1944, p. 73).
The intermediate diameter is the dimension at right angles to the long
axis of the grain, as seen in a microscope slide the cover slip of which
has been pressed firmly down during mounting (in which case the grains
come to lie with their shortest axis vertical and their longest and inter-
mediate axes in the plane of the slide). It is a particularly useful
measure of size, since it is the same dimension as is estimated by sieving.

V. MINERALOGICAL COMPOSITION OF THE HEAVY MINERAL
BEACH SAND CONCENTRATES.

The weight percentage of heavy minerals in the samples of natural
concentrate ranges up to 95.2%, and in most cases it is greater than 50%
(Table I). The light mineral fraction was composed almost entirely of
quartz grains ; very little felspar is present.

In the accompanying Table the numbers indicate weight percentages,
and the symbol “ x ” that the mineral is present in amounts less than
0.1%. The more common species are on the left, and the rare ones to
the right. For convenience, the degree of abrasion of the sands,
calculated as the number percentage of rounded zircon grains in each
sample, is incorporated. The following abbreviations are used : —

Table III. — Mineral Analyses of Heavy Mineral Samples in Percentages by Weight, and Abrasion Grain Number Percentages.

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND

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Table III. — continued.

MINERAL BEACH SANDS OF SOUTHERN QUEENSLAND, PART II

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PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND

Zir, Zircon ; Ru, Rutile ; II, Ilmenite ; Gar, Garnet ; Tour, Tour-
maline ; Mon, Monazite ; Leu +, Leucoxene plus some limonite ; Chr,
Chromite ; Cass, Cassiterite ; Epi, Epidote ; Spin, Spinel ; Mag,
Magnetite ; Cor, Corundum ; Hyp, Hypersthene ; Andal, Andalusite ,
Horn, Hornblende ; Sph, Sphene ; St, Staurolite ; Ky, Kyanite ; Ab;
Abrasion (Grain Number % .Rounded Zircons).

VI. DESCRIPTION OF THE MINERALS.

Zircon. — The zircon grains usually are rounded, the percentage of
such grains in the samples ranging from 45 to 81%. The remainder
are euhedral and subhedral. None is angular from fracture. The
euhedral grains commonly are long tetragonal prisms capped at both
ends by pyramids (Plate III, figure 1). Short or stumpy prisms capped
by pyramids are rarer. The grains are usually clear ahd colourless,
only occasionally brownish. Their surface often shows a high polish.
Inclusions are not abundant ; they are rarer in the rounded than in the
euhedral and subhedral grains. This may be because inclusions are
points of weakness in the crystals, and abrasion liberates them. Inclusions
are usually minute, and, when examined with a high-power magnification,
most of them are seen to be irregular-shaped, hollow cavities (probably
gas-filled) and dust-like spots, but a few are crystals of rutile, iron ore,
monazite, zircon and apatite. Zoning has been observed in a few pf
the grains, but the number is very small in all the samples. The specific
gravity of the zircon from Tugun beach sand is 4.68. The results of
sieve analysis of three widely separated samples of zircon concentrate
are shown below.

Loc.

Size of Openings in Millimetres

0.251-.178

.178-. 152

.152-. 124

.124-. 104

.104-. 076

< .076

Byron Bay

Cudgen

Tugun

0.3%

1.6

0.5

9-0%

8.0

2.4

3.8%

8.1

4.4

63.0%

67.5

73.9

18.4%

11.8

14.9

5-5%

2.9

3.8

The median diameter of the Byron Bay zircon is 0.112 mm., the
Cudgen zircon 0.113 mm., and the Tugun zircon 0.112 mm.

Rutile. — The rutile grains generally are rounded, although edges of
prism faces often can be seen (Plate III, fig. 2). In colour, the grains
range from yellow-red through foxy-red and deep reddish-brown to
black. Some “ black ’• grains are faintly translucent in places when
examined with a combination of transmitted and reflected light employing
a powerful artificial light and a bull’s-eye condenser. Samples of this
“ black ” rutile submitted to chemical analysis showed that it contains
up to 3.5% Fe O. Under reflected light the grains exhibit a high metallic
to vitreous lustre. Striations are visible on some of the grains. As all
gradations between the above colours have been observed, it is impracti-
cable to divide the rutile into varieties based on colour. The specific
gravity of the rutile from Tugun beach sand is 4.21. The results of
sieve analysis of three samples of rutile concentrate are given below.

MINERAL BEACH SANDS OF SOUTHERN QUEENSLAND, PART II. 79

Loc.

Size of Openings in Millimetres

0.251-.178

.178-. 152

.152-. 124

.124-. 104

.104-. 076

<.076

Byron Bay

3.0%

8.7%

7.8%

70.5%

9.2%

. 0.8%

Cudgen

2.0

8.3

5.9

70.6

12.8

0.3

Tugun

3.8

9.9

7.3

71.7

6.9

0.2

The median diameter of the Byron Bay rutile is 0.113 mm., the
Cudgen rutile 0.114 mm., and the Tugun rutile 0.114 mm. Thus, the
rutile grains are approximately equal in size to the zircon grains.
Qualitatively this was found to be so in all the samples.

Ilmenite. — The ilmenite grains usually are not as well rounded as
the zircon and rutile grains, although ilmenite is slightly softer than
these minerals. Very few of the grains, however, show crystal edges.
In some, the surface presents a pitted appearance with irregular-shaped
cavities. Almost all are fresh and unaltered ; very few show partial
alteration to leucoxene. Under reflected light they appear greyish-black,
sometimes with a purplish sheen. According to Miller (1945, p. 69), an
X-ray examination of the ilmenite carried out by the United States
Geological Survey has shown that it is true ilmenite and not arizonite
(Fe2033Ti02), the so-called “ ilmenite ” of Southern India. The specific
gravity of the ilmenite from Tugun is 4.69. The results of sieve analysis
of four widely separated samples of ilmenite concentrate are shown below.

Size of Openings in Millimetres

Loc.

>.178

.178-. 152

.152-. 124

.124-. 104

. 104-.076

<.076

New Brighton

1-4%

15-2%

6-4%

60.5%

12-7%

3-8%

Tugun

5.2

9.6

7.7

68.8

6.4

2.3

Broadbeach . .

6.4

12.4

10.8

61.5

6.9

2.0

Inskip Point . .

5.2

12.3

9.5

60.6

11.2

1.2

The median diameter of the New Brighton ilmenite is 0.116 mm.,
the Tugun ilmenite 0.117 mm., the Broadbeach ilmenite 0.118 mm., and
the Inskip Point ilmenite 0.117 mm. The above figures indicate that
the ilmenite grains on the whole are only a fraction larger than the
zircon and rutile grains.

Garnet.— The garnet grains are angular and irregular in shape (see
Plate I, fig. 2). Apparently they are fragments broken from larger
crystals, and their angularity, which contrasts with most of the other
minerals, is due to greater hardness. The grains exhibit a sub-conchoidal
fracture and vitreous lustre. The surface is sometimes pitted and etched,
the etching giving an irregular, hackly appearance. Mackie (1923,
p. 147) has referred the occurrence of etched garnets in the sandstones
of Scotland to interstratal solution, and Bramlette (1929, p. 336) has
concluded that the etching is produced by acid or alkaline solutions
circulating through sediments subsequent to transportation and deposi-
tion, and that it is generally an authigenic change. In colour the grains
range from almost colourless to pinkish or violet-red, the most common
colour being a pale watery-pink. Inclusions are not abundant ; some
have been identified as quartz, iron ore, zircon and rutile. The refractive
index of the garnet from Tugun beach sand is very close to 1.80, and

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND

the specific gravity is 4.15. The garnet was identified as the variety
almandine, which Milner (1940, p. 233) says “ is the most widespread
and persistent variety of garnet in detrital sediments.” It is moderately
magnetic. The results of a sieve analysis of a garnet concentrate from
Tugun are shown below.

Loc.

Size of Openings in Millimetres

>.251

.251-. 178

.178-. 152

.152-. 124

.124-. 104

<.104

Tugun

2.5%

49.5%

25.9%

9-3%

10.8%

—

The median diameter of this garnet is 0.179 mm. The grains are
much larger than the zircon, rutile and ilmenite grains. Qualitatively
this was seen in all the samples, and no mineral grains larger than garnet
occur in the heavy mineral concentrates.

Tourmaline. — The tourmaline grains are usually rounded, and some-
times have a high degree of sphericity. Prismatic crystals with rounded
ends, some showing vertical striations, are also present. The most
common colours are pale smoky-brown and pale smoky-grey. Occasionally
they are yellow, blue, green and parti-coloured. Although no quanti-
tative determination of size was carried out, most of the grains were
seen to be larger than the zircon, rutile and ilmenite (Plate II, fig. 1).
Inclusions are not common ; they are usually bubbles and long needle-
like particles without definite orientation. The round tourmaline grains
are easily recognised by the comparatively low refractive index, low
double refraction, and the strong pleochroism. In the subhedral grains
the absorption o> > e is apparent. The prismatic grains show straight
extinct ion.

Monazite. — The monazite grains are usually well rounded (Plate III,
fig. 3), but occasionally subhedral grains occur. The grains are pale
honey-yellow in colour, and have a resinous lustre. Inclusions are rare,
and where present they appear mainly to be gas-filled cavities. The
pacific gravity of the monazite from Tugun beach sand is 5.19. A sieve
analysis gave the following result :

Loc.

Size of Openings in Millimetres

>.124

.124-. 104

. 104-.076

< .076

Tugun

—

5.5%

72.7%

, 91 7 0/

41. t /0

A cumulative frequency curve was constructed from the above
information, and the median diameter was 0.086 mm. Thus, the monazite
grains are fairly uniformly small, and usually smaller than most of the
other minerals. Probably this results from the softness of monazite.
A chemical analysis of a sample of monazite concentrate (98% pure)
from Burleigh heavy mineral beach sand contained 7.1% of thoria.

Leucoxene. — The grains of leucoxene are dull-white to yellow-white
when examined by reflected light, often having the aspect of unglazed
porcelain with a rough, pitted surface. They are usually well rounded,
and are non-magnetic. Leucoxene (mainly hydrated titanium dioxide)

MINERAL BEACH SANDS OF SOUTHERN QUEENSLAND, PART II. 81

is an alteration product of titanium-bearing minerals and, according to
Tyler and Marsden (1938, p. 55), it “ develops from the surface weathering
or hydrothermal alteration of sphene or ilmenite.” The grains are
approximately equal in size to the zircon and rutile grains in the
concentrates. .

Cassiterite. — The cassiterite grains are often subhedral or angular,
and are the least well rounded of the heavy minerals with the exception
of garnet (Plate III, fig. 4). The grains are variable in colour ; they are
pale grey, orange, and reddish-brown. This variability is due to traces
of tantalum and niobium. The colour is often irregularly distributed in
the grains, and zoning is not uncommon. These characteristics were
very useful in the recognition of the mineral. The specific gravity of
Tugun cassiterite is 6.90. The results of sieve analysis of two widely
separated samples are shown below.

.Size of Openings in Millimetres

Loc.

0.124-.104

0. 104-.076

<0.076

Byron Bay

5.8%

80.1%

14.1%

Tugun

0.1

19.5

80.4

The median diameter of the Byron Bay cassiterite is 0.084 mm.,
and Tugun less than 0.076 mm. The cassiterite grains are usually
smaller than all the other minerals.

Chromite. — The chromite grains are usually rounded, but occasional
examples retain the characteristic octahedral form with only slight
rounding at the angles. Some are opaque ; others are almost completely
translucent. In contrast to brownish-black, opaque interiors, a number
has greenish-brown margins which are isotropic. Under reflected light
the grains exhibit a submetallic lustre, less brilliant than that of ilmenite.
They are approximately equal in size to the zircon and rutile in the
concentrates.

Epidote. — The epidote grains are yellowish-green, often cloudy from
alteration. While some are rounded, others are prismatic and angular.
They are slightly pleochroic, from pale lemon green to green, biaxial and
negative, with high refractive index and high birefringence. The grains
are usually slightly larger than those of zircon and rutile in the con-
centrates ; apparently this is related to the lower specific gravity of
epidote (3.4).

Spinel. — The spinel grains are usually dark green or greenish-brown
in colour ; some are bright green. They are usually rounded, but traces
of the octahedral habit can be seen. This rather rare mineral is recognized
by its non-magnetic or weakly magnetic properties (in contrast to the
moderately magnetic garnet), and by its colour and isotropism.

Magnetite. — Very few octahedral crystals are present ; almost all are
irregularly shaped. They are black, and some are altered to reddish-brown
limonite. The lustre is silver-grey in reflected light.

Corundum. — The sparse corundum grains are blue, and are referred
to sapphire. The colour is often unevenly distributed in the grains
which are faintly pleochroic and optically uniaxial and negative. The
refractive index is high, and birefringence is weak.

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND

Hypersthene. — The rare grains of hypersthene are pale brownish-
green, and the prismatic habit and distinct cleavage usually is apparent.
They display characteristic pleochroism from pale green to pale pink.
Inclusions of iron ore are present in some.

Andalusite. — The sparse grains of andalusite are pale pink, and
occur as subhedral and rounded particles usually somewhat larger than
the zircon, rutile and ilmenite. This apparently is related to the specific
gravity (3.1), and hardness (7.5) of andalusite. The double refraction
is low, and they are pleochroic from pale pink to colourless. Some
grains are cloudy from alteration.

Hornblende. — This mineral occurs as green, fibrous, flaky grains
with pleochroic absorption parallel to the horizontal crosswire. The
interference colours are of low order, and the extinction angle is 13
degrees. There is good cleavage parallel to the length of the flakes.

Sphene.- — The sparse grains of sphene are of characteristic brown
colour, often somewhat clouded through decomposition to leucoxene.
They are irregularly shaped, and not well rounded. The refractive index
is very high, total extinction is absent, and the interference colours
usually show a bluish tint.

Staurolite.— The staurolite grains are brown or reddish-brown in
colour, and usually rounded,. Some show good cleavage, and exhibit
strong pleochroism, ranging from pale reddish-brown to dark brown.
These properties serve for their identification. Inclusions occur, most of
them appearing to be black iron ore and quartz.

Kyanite— The very few grains of this mineral are elongate, subhedral,
and rounded at the ends. They are almost colourless. Good cleavage
is apparent, and some minute inclusions were seen.

Quartz.- — Although the quartz grains were not examined in all the
samples, the light fraction from some of them was mounted in Sirax
(refractive index 1.80) And examined under the microscope. Generally
they are larger than the heavy minerals, and usually subangular. Some
of the grains are water-clear, but most are cloudy with minute inclusions
or slightly affected by yellow iron staining. Many of the inclusions
appear to be gaseous and fluid, but some prisms of tourmaline, needles
of rutile, and particles of iron ore were recognised. Some of the grains
exhibit strain polarisation shadows.

Abrasion.

The heavy minerals are fairly well abraded. This is apparent under
the microscope even without a quantitative determination of their degree
of abrasion. The results of the quantitative determination (Table III)
show that there is no general trend of change in the degree of abrasion
from south to north along the coast. However, the grain number per-
centage of rounded zircons usually is less in the samples from near the
major coastline breaks, particularly to the north of them, and from
about the headlands of Mesozoic sandstone. Thus, the abrasion per-
centages for samples 14, 15 and 16 close to the Nerang River mouth
are respectively 70%, 69% and 61%, while that of sarqple 13 to the
south, is 78%, and that of sample 17 to the north, is 80%. Similarly,
the grain number percentage of rounded zircons in sample 40 from near
Alexandra Headland is as low as 45%, but this decrease in abrasion
again is only local.

MINERAL BEACH SANDS OF SOUTHERN QUEENSLAND, PART II.

It is of interest to note that the sand from the bore sunk with casing
to 36 feet in the Eighteen Mile Swamp on North Stradbroke Island
contains only 53% rounded zircons, while the samples from the beach
and dunes in the neighbourhood all contain a much higher grain number
percentage of rounded zircons. The sand from this cased bore has come
mainly from below present sea-level, and the smaller degree of abrasion
probably results from it not having been subjected as much to the action
of waves and wind as the beach and dune sands. The number percentage
of- rounded zircon grains in the 50 samples ranges from a minimum of
45% to a maximum of 81%.

That there is no general decrease or increase in abrasion from south
to north suggests that the heavy minerals have not come solely from
one locality in the extreme south or north. Indeed, the definite local
trends which appear to be related to major coastline breaks and sand-
stone headlands, suggest that additions of heavy minerals have been
made to the shore at various places along the coast.

VII. GEOGRAPHIC DISTRIBUTION OF THE
HEAVY MINERALS.

The Table of Mineral Analyses shows that the heavy mineral beach
sand concentrates are composed essentially of zircon, rutile and ilmenite.
All the other minerals together usually make up less than 5% of the
total heaVy mineral weight percentage. However, there are large changes
in the individual weight percentages of zircon, rutile and ilmenite in
the concentrates from south to north along the cohst.

In the area to the south of the Nerang River zircon is the most
abundant mineral, usually with rutile in second place and ilmenite third.
However, a gradual decrease in the percentage of zircon and a slight
increase in the percentages of rutile and ilmenite from south to north
is noticeable in this area.

On South and North Stradbroke Islands the weight percentages of
zircon, rutile and ilmenite in the concentrates are approximately equal,
except for a slight decrease in the zircon and an increase in the ilmenite
content northwards. Throughout this area, the rutile content remains
almost stationary. Therefore, on Stradbroke Island the zircon content
usually is lower, and the rutile and ilmenite contents slightly higher,
than in the concentrates from south of the Nerang River mouth.

On Moreton Island and in all places further north ilmenite is the
predominant heavy mineral. It forms over 50% of the heavy minerals
in all the samples examined, and the weight percentages of zircon and
rutile are approximately equal. In the stretch of coastline northwards
from the South Passage, the weight percentages of zircon and rutile
generally are each less than 25%, with rutile slightly less than zircon.
Throughout this area the most striking trend in the distribution of the
heavy minerals is the decrease in zircon content and the increase in
ilmenite northwards.

The distribution of the other minerals is rather erratic, but several
definite features are evident. The most conspicuous is the much higher
garnet content in the neighbourhood of the headlands of Mesozoic sand-
stone which occur at intervals along the coast from Cape Moreton north-
wards. To the south the garnet content is not greater than 1.4% in
any of the samples studied. Northwards, however, the garnet content
is as high as 5.2% in the Caloundra Head beach sand, 8.3% at Alexandra

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND

Headland, 7.1% at Pt. Arkwright, 3.5% at Paradise Caves (Coolum
Beach), 1.5% at Noosa Head, and 2.1% in the sample from 3 miles N.
of the Noosa River mouth. Northwards from Double Island Point,
where there are no further outcrops of Mesozoic sandstone, the garnet
content is again low, less than 1% in the samples studied.

Another feature is the much higher tourmaline content compared
with garnet in the samples from Stradbroke Island. Elsewhere the
garnet content is usually slightly greater than tourmaline ; on Stradbroke
Island the tourmaline content is generally more than 1.5%, while the
garnet is less than 0.5% and is as low as 0.1% in one sample. This may
result from the greater stability of tourmaline. On Stradbroke Island
no possible source rocks outcrop along the coast, and the heavy minerals
appear to have had a long history since liberation.

Monazite, like garnet, is usually more abundant in the vicinity of
outcrops of Mesozoic sandstone along the coast. For example, it is as
much as 2.4% in the Caloundra Head sample, 1.1% at Alexandra Head-
land, 1.5% at Pt. Arkwright, 1.1% at Paradise Caves (Coolum Beach),
1.0% at Noosa Head, 1.2% in the sample from just N. of Noosa, and
1.4% in the Double Island Point sample. In almost all other localities
the monazite content is less than 1%, and on North Stradbroke Island
it is usually less than 0.5%.

The Table of Mineral Analyses shows that epidote is often present
in samples in which the tourmaline content is higher than usual. This
association apparently is related to the similar, low specific gravity of
these two minerals. Although chromite is irregular in its abundance, it
shows a drop in quantity in the Stradbroke Island samples. The distribu-
tion of other minerals is irregular. Very few grains of corundum,
hypersthene, andalusite, hornblende, sphene, staurolite and kyanite are
present in any of the samples.

VIII. ORIGIN OF THE HEAVY MINERALS.

Samples from outcrops of possible source-rocks in south-eastern
Queensland and northern New South Wales were examined in the hope
of obtaining evidence concerning the origin of the heavy minerals. They
were broken down in the manner described above, and their heavy
mineral assemblages studied under the microscope.

River sand concentrates also were examined from streams entering
the sea between Yamba in northern New South Wales and Caloundra
in southern Queensland. Samples were panned from bars and banks,
and from the shallow, upper reaches of channels and the beds of streams.

Geology of the Region.

The following brief account of the geology of the region has been
prepared from a number of papers, and from the held observations of
the writer. For the sake of clarity, Queensland rivers have been omitted
from the geological map (Text-hg. 5).

The basement rocks outcrop along the coast in the southern part
of the region. They make up, in some places with Tertiary basalt, all
the headlands as far north as South Nobby Headland. They are low-rank
metamorphics (greywackes, slates, quartzites, jaspers and phyllites) of
Lower Palaeozoic age, which were originally laid down as sediments in
the Tasman Geosyncline. From the South Coast (New South Wales

AL BEACH SANDS OF SOUTHERN QUEENSLAND, PART II.

TEXT- FIGURE 5. — GEOLOGICAL MAP OF
REGION, SHOWING PLACE
OF COLLECTION OF ROCK
SAMPLES STUDIED.

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND

border to Southport) area of Queensland northwards, these rocks trend
NNW. away from the coast. In SE. Queensland they are known
collectively as the Brisbane Schists, and the name Neranleigh Series has
been given to the subdivision consisting largely of greywackes.

Further inland, in the southern half of the area, there is a small
outcrop of Devonian basic volcanics, chert and limestone, and there are
some areas of Permian sediments, the largest of which is in the Drake-
Boorook district. In the northern half of the area, away from the coast,
there are also some small areas of Permian sediments which are mainly
of shallow-water marine origin, but near Warwick, freshwater deposits
are interbedded.

In late Permian times, these Palaeozoic metamorphic and
sedimentary rocks were intruded by numerous granitic masses.
By far the largest of these is the New England-Stanthorpe com-
posite batholith. It is centred about Tenterfield and covers an
area of several thousand square miles. Andrews (1907) has recognised
a number of phase-types in this mass: — namely, “ Grey Felspar
Porphyries/’ “ Blue Granite,” “ Sphene Granite Porphyry,” and
“ Coarse Acid or Tin Granite ” — the main intrusions becoming pro-
gressively more acid and having their maximum development in the
northern part of the mass. An extensive development of greisen and
pegmatite occurs about the peripheries of these acid intrusions, and
with them are associated important ore deposits containing tin, bismuth,
tungsten and molybdenum, as well as monazite, gold and other valuable
minerals (Andrews, 1905 a ; 1905 b). The granitic masses further north
are similar but smaller in area (Bryan, 1922, pp. 148-157). Probably
some of them were comagmatic. At. least two of these granitic masses
(near Kingaroy and Mount Perry) are known to have small deposits of
rutile and ilmenite around their margins. According to Cribb (1943,
p. 39) the Kingaroy mass, is a medium-grained biotite granite, and in
the neighbourhood of the deposits it gradually assumes a gneissic
structure. “ The granite is intruded by dykes of pegmatite and aplite,
usually of small size. Black tourmaline is abundant in the former in
association with white felspar, quartz, and muscovite.” The rutile is
seen now mainly as shoad material, “ occurring in grains and pieces, the
larger sizes usually with an elongated outline up. to 1J inches long, and
generally exhibiting vestiges of crystal faces with rounded angles.
Forms present include pyramids and 1st and 2nd order prisms.” Some
ilmenite is associated with the reddish-brown rutile, and both are found
as sparsely scattered nodules surrounded by thin segregations of biotite
in the granite. Fisher (1945) states that the rutile near Mount Perry
occurs in small veins less than 8 inches wide associated with finely
granular quartz in the granite. According to Morton (1946), small
rutile lode deposits also are found associated with Permian pegmatites
in North Queensland. Around the granitic masses, contact metamorphic
rocks such as garnet hornfels and spotted slate are often present.

Most of these late Permian granitic bodies must have extended to
very near the surface, as they were unroofed in at least Middle Triassic
times. Their injection was associated with a major movement of uplift,
accompanied by folding and mountain building. The Palaeozoic Tasman
Geosyncline was finally drained, and freshwater lakes were left in its
place. The position of these lakes in Upper Triassic and Jurassic times
marginal to the large New England-Stanthorpe granitic mass, and the

MINERAL BEACH SANDS OF SOUTHERN QUEENSLAND, PART II.

other granitic masses in the area, is proof that relatively high land
existed there. In places the lakes extended some distance east of the
present coastline, and it is possible that granitic masses also were
associated with high land along an eastern margin now beneath the
Pacific Ocean. Earth-movements after the close of the Jurassic drained
the lakes, and the rocks of the area in places were considerably folded.

A large part of this coastal area is -covered by these Mesozoic fresh-
water sediments. In Queensland the Upper Triassic lacustrine sediments
are known as the Ipswich and Bundamba Series, while those of Jurassic
age are the Walloon Series. In northern New South Wales the equiva-
lents of the Bundamba and Walloon Series are known as the Clarence
Series. Near the margins of both the Walloon and the Clarence Series,
these Jurassic sediments can be seen resting on well-eroded, Permian
granitic surfaces. In the Maryborough-Bundaberg district, there is a
small belt of shallow-water, marine and freshwater sediments of Cretaceous
age. At Point Arkwright and Noosa Head the Jurassic sandstones have
been intruded by very small masses of granodiorite of late Mesozoic age.

In many places these Palaeozoic and Mesozoic rocks are overlain by
Tertiary volcanics. These are mainly basic in composition (andesites
and basalts), and are principally Pliocene in age. They are particularly
abundant in the region about the Queensland-New South Wales border,
and formerly covered much of the New England Tableland. They out-
crop along the coast at such places as Fingal Head, Point Danger,
Burleigh Head, Double Island Point, and Indian Head.

The Pliocene basalts were deeply dissected following a major move-
ment of uplift at the close of the Tertiary, and the Mesozoic and Palaeozoic
rocks which underlay them were exposed again in many places. The
forces of erosion were so active that the basalts were removed almost
completely from large areas, and occur only as isolated, residual cappings
at the present time.

The streams traversing these rocks are quick-flowing in their upper
reaches, but except in times of flood, they are rather sluggish in their
lower reaches. Stream-deposits of economic value occur. Alluvial tin
is worked in the upper reaches of the Clarence River, and zircon, monazite,
topaz, garnet, spinel and sapphires are recovered, as well as some gold
and platinum.

Pleistocene and Recent beach and dune sands cover much of the
country fringing the present coastline. The large coastal islands are
almost entirely covered with sand-dunes, and only at a few places close
to sea-level do rocks outcrop.

Nature and Places of Collection of Rock and
River Sand Samples.

The location of tfye places where the rock samples and river sands
U and V were collected is shown in Text-figure 5, and those of the other
river sands in Text-figure 1.

Rock Samples.

A. Greywacke from Neranleigh Series exposure in main road in cutting,

Burleigh Head.

B. Greywacke from exposure in railway cutting, \\ miles S. of Nerang

railway station. In type area of Neranleigh Series.

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND

C. Greywacke from Neranleigh Series exposure at Brookfield, near
Brisbane.

D. Coarse-grained, acid New England granite from outcrop alongside
New England Highway, N. of Tenterfield.

E. Pegmatite near periphery of New England batholith, NE. of Tenter-
field.

F. “ Basic ” blue granite from outcrop, 2 miles E. of Tenterfield.

G. Sphene granite-porphyry from outcrop near Wallangarra.

H. Coarse-grained Permian granite from Stanthorpe.

I. Permian granite from Yarraman, S. of Nanango.

J. Late Cretaceous granodiorite intruding Jurassic sandstone at Noosa
Heads.

K. Tertiary andesitic-basalt capping Ordovician greywacke at Burleigh
Head.

L. Tertiary basalt from Indian Head.

M. Lower Jurassic sandstone from outcrop near Yamba.

N. Clarence Series sandstone from exposure in road cutting near
Tabulam.

O. Lower Jurassic Clarence Series sandstone (slightly weathered) from
exposure in road cutting near Broken Head, about 4 miles S. of
Byron Bay.

P. Upper Triassic Bundamba Series sandstone from outcrop near
Beaudesert, in Logan River valley.

Q. Upper Triassic sandstone from Cape Moreton.

R. Upper Triassic Bundamba Series sandstone from Caloundra Head.

S. Jurassic sandstone from Alexandra Headland.

T. Jurassic sandstone from Point Arkwright.

River Sand Samples.

U. Oraro River, just W. of Grafton. The Oraro River entirely traverses
Clarence Series strata.

V. Mann River, on Glen Innes-Grafton road 78 \ miles from Grafton
within the outcrop of the New England granitic mass.

W. Bluff River, between Tenterfield and Glen Innes in the heart of
New England granitic mass.

X. Sandy Creek, a tributary of the Clarence River, between Tabulam
and Tenterfield. On New England granite, not far from the eastern
edge of the batholith.

Y. Plumbago Creek at Drake in the centre of a small block of Permian
sediments, immediately E. of New England granitic mass.

Z. Tributary of the Richmond River traversing area of Clarence Series
strata, between Tabulam and Casino. Relatively large quantities
of heavy mineral concentrate were obtained at each panning.

ZA. Richmond River at Casino. The concentrate was obtained by
panning river sand resting on Clarence Series sandstone. Tertiary
basalt is very abundant in the vicinity.

ZB. Tweed River near Uki. The Lower Palaeozoic metamorphics are
intruded by a small Permian plutonic mass and overlain in many
places by Tertiary volcanics in the region.

MINERAL BEACH SANDS OF SOUTHERN QUEENSLAND, PART II. 89

ZC. Currumbin Creek, which flows through Lower Palaeozoic greywackes
and slates capped by Tertiary basalts. Only a small quantity of
heavy mineral concentrates was obtained at each panning.

ZD. Brisbane River at Moggill. In an area of Upper Triassic sandstones
and Lower Palaeozoic metamorphics intruded by some Permian
granitic masses.

ZE. Stanley River at Somerset Dam. Rocks mainly andesitic material,
but some plutonic rocks of intermediate composition present.

Analyses of Heavy Mineral Fractions of the
Rock Samples.

In the accompanying Table the minerals are arranged in order
similar to Table III and the same abbreviations are used, the symbol
“ x ” indicating that the mineral is present in amounts less than 0.1%.
For convenience, the degree of mineral abrasion in the met amorphic
and sedimentary rocks, calculated as the grain number percentage of
rounded zircons, is incorporated in this Table. Additional abbreviations
used are

I.F., Index-figure ; II +, Ilmenite plus chromite when present ;
Pyrox, Pyroxene ; Horn+, Hornblende plus some chlorite ; Ap, Apatite ;
Bio-f , Biotite plus some chlorite ; Py, Pyrites ; To, Topaz ; Ana, Anatase.

Discussion.

[a). Greywackes.

The index-figures in the preceding Table show that the weight
percentage of heavy minerals in the greywacke samples is exceedingly
small, less than 0.01%. The grain size is smaller than that of any of
the beach sands. In sample A the mean of the zircon intermediate
diameters was 0.063 mm., the largest 0.086 mm., and the smallest
0.049 mm. In sample B the mean was 0.064 mm., the largest 0.098 mm.,
and the smallest 0.036 mm., while in sample C the mean was only 0.055
mm., the largest 0.061 mm., and the smallest 0.049 mm. The means of
100 intermediate diameters each of the beach sand zircon from Byron
Bay, Cudgen and Tugun were 0.114 mm., 0.115 mm., and 0.114 mm.
respectively, which is practically the same as the median diameters. The
zircon size in the greywacke samples is therefore considerably smaller
than in the beach sands (Plate III, fig. 1, and Plate V, fig. 1). The degree
of zircon abrasion (Table IV) usually is not as great as in the beach
sand concentrates.

The heavy mineral assemblages of the greywackes are made up
essentially of zircon, ilmenite, leucoxene and tourmaline. All the other
minerals, such as rutile, garnet, cassiterite, epidote, spinel, kyanite and
.anatase are very scarce, and together form less than about 5%.

These results, particularly the exceedingly small heavy mineral
content, the small grain size, the heavy mineral composition, and certain
mineralogical properties such as the large number of dusky zircon grains,
indicate that the greywackes have not played a major role as a con-
tributor of the heavy minerals in the beach sand deposits.

(b). Granitic Rocks.

The index-figures in Table IV show that the weight percentage of
heavy minerals in some of the granitic samples is comparatively large.
This is mainly because the ferromagnesian silicate minerals, which have

Table IV.— Mineral Analyses of Heavy Mineral Fractions of the Rock Samples, and Abrasion Grain Number Percentages.

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND

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MINERAL BEACH SANDS OF SOUTHERN QUEENSLAND, PART II. 91

specific gravities greater than 2.86, are included. These ferromagne$ian
silicates are the heavy essential minerals, as opposed to the heavy
accessory minerals. Becausfe of the influence of these minerals upon the
index-figures, it is natural that the basic “ granite ” and granodiorite
samples have the highest index-figures. There is a wide range in the
index-figures of the five samples from the New England batholith,
apparently as a result of contamination by assimilation of foreign material,
and differentiation of the mass.

The means of 100 zircon intermediate diameters in the seven samples
are : — Sample D, 0.115 mm.; E, 0.112 mm.; F, 0.125 mm.; G, 0.105 mm.;
H, 0.114 mm.; I, 0.110 mm.; J, 0.100 mm.

Thus, the zircon in the six Permian granitic samples is either slightly
larger (sample F), or almost exactly the same size (samples D, E, G,
H and I) as the beach sand zircon. The sparse zircon in the Cretaceous
granodiorite (sample J) is smaller.

The more abundant heavy accessory minerals in the seven granitic
rocks are zircon, apatite, sphene, magnetite and ilmenite. The heavy
essential minerals are biotite and hornblende. In four of the rocks
(the more acidic), zircon is by far the most abundant mineral in the
non-magnetic, heavy mineral fraction. That it is seldom seen in thin
sections apparently is due partly to the very small area of such sections,
and partly to the fact that it is prone to break away during the grinding
preparations. It is an early product of crystallization and is often
surrounded by an air pocket.

In three of the rocks apatite is the most abundant non-magnetic
heavy mineral. Sphene is common in the sphene granite-porphyry, and
is visible even in the hand specimen. In four of the other granitic rocks
also it was found, but very scarce. Rutile is not abundant, but occurs
in five samples. In two samples, magnetite is hot in excess of ilrpenite.
While pneumatolytic minerals are not abundant, tourmaline, cassiterite,
monazite, topaz, and anatase are present in the Permian samples. They
occur more in the acidic samples, tourmaline and monazite being the
most widespread. The presence of pink garnet (almandine) in sample H
from Stanthorpe is suggestive of rock assimilation and contamination
near the margin of the batholith.

The geology of the region indicates that the most feasible major
primary source for the beach sand heavy minerals are the Permian
granitic rocks. The fact that zircon of very similar size and appearance
to the beach sand zircon is common in the six Permian granitic samples,
and that rutile, ilmenite, magnetite, monazite, tourmaline, cassiterite,
epidote, hornblende, sphene, and some pink garnet are also present is
confirmatory evidence. The two very small outcrops of late Cretaceous
granodiorite, at Point Arkwright and nearby Noosa, could not have
played any major role as contributors of the beach sand heavy minerals.
The very sparse zircon in the granodiorite from near Point Arkwright
(sample J) is of a smaller size than the beach sand zircon, while more
of the grains have a cloudy or dusky appearance. Pneumatolytic
minerals such as tourmaline, monazite, topaz, cassiterite and anatase
are also lacking from the heavy mineral assemblage.

(c). Basalts.

As it was realised that basic volcanic rocks could not be important
source rocks jfor most of the heavy minerals in the beach sands, only
two samples of Tertiary basalt from the area were chosen for examination.

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND

They were the Burleigh Head andesitic basalt (sample K) and the Indian
Head basalt (sample L). Table IV shows that the index-figures for
these rocks are high. This is chiefly because of the presence of a great
quantity of the heavy essential mineral, augite. It is the most abundant
heavy mineral in both samples. Richards (1916, p. 174) examined thin
sections of the Burleigh basalt and stated that “ the augite occurs in
sub-ophitic rounded patches but occasionally in long patches 1 mm. in
length.” Olivine is not present.

The heavy accessory minerals in both samples are mostly black
iron-ore, although some apatite is present. Treatment with the electro-
magnet showed that the bulk of the iron-ore material is moderately
magnetic, and it has been referred to ilmenite, although a very small
amount of chromite may be present. A small amount of the material
is highly magnetic, and this has been referred to magnetite. In thin
sections of these basalts the iron ore has been found to occur in granules
and in rods, the latter averaging 0.4 mm. in length. It will be recalled
that Tertiary volcanics, particularly those of basic composition, are very
extensively developed in certain parts of the area, while evidence indicates
that they formerly covered a much greater area than at present. From
their distribution, and the fact that ilmenite is common in the samples,
it is apparent that they have contributed at least some of the ilmenite
in the beach sands, and it is possible that they may be an important
source rock for this mineral.

(d). Sandstones.

The weight percentage of heavy minerals in the eight samples of
Mesozoic freshwater sandstone varies from less than 0.1% to as much
as 1.5%, and is highest in the Clarence Series sandstones. In the Upper
Triassic Bundamba sandstones and the Jurassic sandstones from southern
Queensland the weight percentage of heavy minerals is 0.1% or less.
This is comparable in amount with that of the freshwater Narrabeen
sandstones from near Sydney, studied by Culey (1932). Since the weight
percentages of heavy minerals in the three Clarence Series samples are
0.7%, 0.8% and 1.5%, it would seem that these freshwater sandstones
are particularly rich in heavy minerals.

Unfortunately, it is not possible at present to give the exact age
and position in the stratigraphic column of the sandstone samples studied.
This is because of the sporadic and often poor nature of the sandstone
outcrops, the lithological similarity of most of the sandstones, the paucity
of exposures containing fossiliferous shale bands, and the scarcity of
detailed geological mapping in the region.

Even without a quantitative determination of size, it is apparent
that the grain size of the heavy minerals generally is slightly larger than
those of the beach sands. The means of 100 zircon intermediate diameters
in the eight samples were found to be : — Sample M, 0.110 mm.; N, 0.152
mm.; O, 0.120 mm.; P, 0.115 mm.; Q, 0.125 mm.; R, 0.120 mm.; S,
0.126 mm.; T, 0.130 mm.

Thus, the zircon usually is slightly larger than the beach sand zircon,
although in samples M and P it is of about the same size. Microscopic
examination of the heavy mineral assemblages has shown that the
minerals are not as rounded as those of the beach sand deposits. The
degree of zircon abrasion (Table IV) is clearly less. It will be noticed
that the sandstone samples collected from near beach concentrates have

MINERAL BEACH SANDS OF SOUTHERN QUEENSLAND, PART II. 93

lower grain number percentages of rounded zircons than the adjacent
concentrates. The heavy minerals usually do not appear to be as well
sorted as those of the beach sand concentrates (see Plate V, figs. 3 and 4).

The heavy mineral assemblages of the sandstones are made up
essentially of the same suite of minerals as the beach sand concentrates,
but the proportions are somewhat different. Although the weight
percentage of zircon in six of the samples is greater than 20%, it is the
predominant heavy mineral in only samples M and Q. Rutile is the
most abundant heavy mineral in samples N and T. In N the content
is as much as 37.9%, and in T it is as high as 40.5%. The iron-ore
content ranges from 2.8% to 22.4%, and it is less than 20% in seven
of the eight samples. The opaque iron ores are not the most abundant
heavy minerals in any sample.

Garnet is the most abundant heavy mineral in samples P, R and S,
and its weight percentage is more than 25% in four of the samples.
In the Caloundra Head sample it is 58.8%, and in the Alexandra Head-
land sample it is as much as 85.4%. As already stated, the garnet
content in the beach sand concentrates usually is markedly higher in
the neighbourhood of the outcrops of Mesozoic sandstone, but it is much
less than that in the sandstone assemblages. The percentage of tourmaline
ranges from 1.1% to 3.9%, and it is usually a little greater than in the
beach sand concentrates. The content of monazite is generally greater
than that in the beach sand concentrates. It ranges from 1.7% to as
much as 6.1%.

The percentage of leucoxene (plus some limonite) ranges from 0.4%
to 41%, but it is greater than 10% in only one sample. The content
is generally higher than in the beach sand concentrates, and the weight
percentage of 41.0% in sample O is conspicuously higher. Sample O is
from an exposure in a road cutting, and weathering in situ may be
partly responsible for this high leucoxene percentage. Nevertheless, an
ilmenite content or leucoxene source originally much greater than that
of any of the other sandstones is indicated. The content of epidote is
slightly greater than in the beach sand concentrates.

From the very rich content of heavy minerals in some of these
sandstones, the larger grain size, the less abraded and less well-sorted
nature of the heavy minerals, the presence of essentially the same suite
as in the beach sands, the striking similarity in appearance under the
microscope of the minerals (see below) with those in the beach sands,
and the absence of any other likely major source rock in the vicinity,
it is concluded that the freshwater Mesozoic sandstones are the immediate
source rocks of most of the heavy minerals in the coastal sand deposits.
The differences in the proportions of the various mineral species in the
sandstone heavy assemblages and the beach sand concentrates apparently
are due, in part, to the fact that much of the ilmenite presumably is
derived from the Tertiary basalts of the region. The amount varies.
The differences apparently are due in part also to the varying degree
of stability of the heavy minerals, and to the fact that in the coastal
sands they have been through at least one more cycle of erosion, trans-
portation and deposition. As Pettijohn (1941, pp. 610-625) and Lincoln
and Clarissa Dry den (1946, pp. 91-96) have shown, the resistance of
garnet to weathering is much less than that of zircon, rutile, tourmaline
and monazite. This would explain the much lower percentage of garnet
in the beach sand concentrates than in the nearby sandstones. Similarly,

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND

the greater abundance of zircon in the beach sand concentrates must be
partly due to its chemical and physical stability, which is known to be
extremely great.

Study of River Sand Concentrates.

In most cases, the amount of heavy minerals in the river sands
was small, and in some it was exceedingly small. Any light minerals in
the panned samples were removed with bromoform. Mechanical analyses
of all except samples Y and ZD, which were too small for accurate
sieving, were then carried out in the same way as for the panned beach
sand concentrates. The results are given in Table V, and the quartiles,
medians, and coefficients of sorting in Table VI.

Table V. — Mechanical Analyses of Heavy Mineral Sand

Concentrates.

Size of openings in millimetres

No.

>.251

.251-. 178

. 178-.152

.152-. 124

.124-. 104

T04-.076

< .076

22.2

40.8

14.8

5.3

3.4

12.7

0.7

9.9

27.9

16.5

8.0

12.0

25.4

0.3

—

14.2

16.0

11.8

5.0

52.6

0.4

7.2

40.4

20.4

10.1

0.3

21.5

0.1

3.2

21.6

17.0

10.4

0.9

44.1

2.7

—

5.0

8.8

8.5

2.1

67.4

8.2

90.1

8.7

0.5

0.2

0.1

0.3

0.1

0.6

3.1

5.1

3.0

2.1

66.2

19.9

64.5

28.9

3.5

0.9

0.2

1.9

0.1

TABLE VI. — First and Third Quartiles, Medians, and Coefficients of
Sorting of the Heavy Mineral River Sand Concentrates.

No. Q1 M 03 So

Millimetres

U ...

.250

.200

.158

1.26

V ‘ ...

.208

.160

.104

1.41

W ...

.161

.102

.088

1.35

.218

.174

.134

1.27

.178

.130

.088

1.41

.104

.092

.082

1.13

>.251

b. >'■ >■ - T

TOO

.088

.078

1.13

>.251

.224

Samples ZB and ZE differ from the beach sand concentrates in that
the maximum sieve-fraction is that retained on the B.S.S. 60 sieve
(aperture size 0.251 mm.). This size fraction proved to be mainly
ilmenite. Samples U, V and X have maximum sieve-fraction percentages
in the 0.251 mm. to 0.178 mm. grade size. This is larger than that of
all except three of the beach sand concentrates obtained from sands
adjacent to coastal outcrops of Mesozoic sandstone. The other four
river sand concentrates have maximum sieve-fraction percentages in the
0.104 mm. to 0.076 mm. grade size.

The median diameters of the heavy mineral river sand concentrates
range from a minimum of 0.088 mm., in the Currumbin Creek sample,
to more than 0.251 mm. in samples from the upper reaches of the Tweed

MINERAL BEACH SANDS OF SOUTHERN QUEENSLAND, PART II.

and Brisbane Rivers. These are larger than any of the panned beach
sand medians, but the median of samples from Currumbin Creek, Bluff
River, and Richmond River (ZA) are smaller. The medians of samples
U, V and X from tributaries of the Clarence River, and Z (from the
Richmond River) are within the size range limits of the heavy mineral
beach sand medians, but all are larger than the average. It is of interest
to note that the median of sample X, from near the margin of the New
England granitic mass, is as much as 0.174 mm., while that of sample W,
from the heart of the eroded batholith, is only 0.102 mm.

Coefficients of sorting for only seven of the river sand concentrates
are given. Quartiles could not be determined for two of the samples
as most of their bulk was retained on the coarsest sieve employed. The
coefficients range from 1.13 to 1.41 ; that is, the heavy minerals are
generally not as well sorted as those of the beach sand samples. The
least well-sorted heavy minerals are those of samples V, W and Z, all
from within the outcrop of the New England granitic mass or close
to its margin.

Log skewness calculations were not made, as it was felt that they
would be of little practical value. However, some knowledge of the
skewness of the heavy mineral samples can be obtained from Tables V
and VI.

The mineral analyses are shown in Table VII, together with the
degree of mineral abrasion (calculated as the grain number percentage
of rounded zircons). The abbreviations used are the same as those in
Tables III and IV.

Discussion.

The degree of abrasion of the heavy mineral river sands is usually
much less than that of the beach sands. Table VII shows that, with
the exception of sample U, the grain number percentage of rounded
zircons in all samples is lower than that in the beach concentrates. The
river sands containing the least number of rounded zircon grains are
those which appear to have been derived directly from the weathering '
of Permian granitic rocks. An increase in abrasion in the river sands
away from the granitic outcrops, particularly in the regions covered by
later, freshwater sedimentary rocks is apparent from the grain number
percentages.

The heavy minerals comprise essentially the same species as are
found in the beach sands. However, there are fairly large differences in
the weight percentages of some of the minerals compared with those
of the beach, and in the number of mineral species in some of the river
sands.

In the samples which appear to have been derived from the dis-
integration of the Permian granitic rocks, more or less in situ, the
percentage of magnetite is high. In the other samples, however, the
content is low. The fact that it is rare in the heavy mineral beach
sands may be taken as indicating a greater distance from its primary
source. As well as high magnetite contents, all except one of the samples
from within and near the granitic outcrops contain more than 30% of
ilmenite. Probably some of this ilmenite came from the Tertiary basalts
1 in the area. In other samples, ilmenite is much more abundant than
magnetite. In sample ZC from Currumbin Creek, which drains an area
consisting only of Ordovician greywacke and Tertiary basalt, the ilmenite

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND

C/2 C/2

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CD CM CM CM | CO H |

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XXX X

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T* 10 10 t> GO

cm’ d CO CM d

1 — 1 1 10 VO (M 0

0 1— 1 d d d

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1 1

MINERAL BEACH SANDS OF SOUTHERN QUEENSLAND, PART II. 97

makes up 86.4% of the heavy minerals, and the fact that the grains are
not rounded suggests that it is derived from the basalt and not from
the greywacke.

The contents of hornblende and sphene are much greater in the
sands from within and near the granitic masses than in those from other
places. This, together with the distribution of topaz, is apparently
related to the comparatively low stability of these minerals. The weight
percentages of zircon and rutile, together with those of garnet and
monazite in particular, are greater in the river sands collected from
regions of Mesozoic sandstone than from the granitic areas. This is
apparently due to the gradual removal of less stable minerals such as
magnetite, hornblende, augite, sphene, apatite and topaz.

From the above it is clear that the major rivers which flow chiefly
through regions of Mesozoic sandstone bring to the sea essentially the
same assemblage of heavy minerals as occur in the beach sands, and
that the streams which drain areas where Tertiary basalts are widespread
transport a much larger proportion of ilmenite seawards. The rivers
in the southern part of the area appear to bring down to the coast less
ilmenite than do those to the north. It is unlikely that any great
quantity of the heavy mineral material derived directly from weathering
of the Permian granitic rocks reaches the Pacific Ocean at the present
time.

The mineralogical composition, size-distribution, sorting, and
abrasion of the heavy mineral river sand concentrates, as well as the
appearance of the heavy minerals under the microscope indicate that
the beach sand heavy minerals could have been derived largely from
the rocks in the hinterland and transported to the sea by the present
coastal streams. Most of the coastal streams are rather slow-flowing in
their lower reaches, but their rate of flow increases considerably following
periods of heavy rainfall, and they are subject to severe floods. Large
quantities of detrital material are then carried to the Pacific Ocean.
Even under normal conditions, tidal currents help to carry some sands
from the lower reaches of the rivers out to sea.

Although the coastal rivers may not always have occupied their
present courses, Craft (1933) considered that the positions of the major
streams were approximately as at present at least before the outpourings
of basic lava in the Pliocene. The rate of flow would be great after
movements of uplift and rejuvenation which occurred in Tertiary times,
and vast amounts of heavy mineral material would be brought down to
the sea and distributed along the coast.

Descriptions of the Heavy Minerals in the Rock and
River Sand Samples.

Zircon.

Greywackes (Plate V, fig. 1). Apart from the small size, the most
distinctive feature is the relatively large number of dusky or cloudy
grains. Unlike the beach sand zircon, many of the grains are crowded
with inclusions which give them this cloudy appearance, and a large
number of them show zoning.

Granitic Rocks (Plate V, fig. 2). Most of the zircons are long
tetragonal prisms capped at both ends by pyramids. A small number
is corroded, and some are well rounded and have a “ waterworn ”
appearance. While these may be xenocrysts, it is possible that they

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND

may all be the result of magmatic chemical corrosion subsequent to
crystallization. Almost all are colourless, but a few crystals are pale
brownish. Inclusions are not common ; a few are crowded with them,
and have a cloudy appearance contrasting with the comparative clarity
of the others. Some of the inclusions are black iron ore (probably
ilmenite), rutile, apatite, and minute zircons, but many are irregularly-
shaped cavities and dust-like spots. A small number of the crystals
show zoning. This is more common in those which have a cloudy
appearance.

Sandstones (Plate V, figs. 3 and 4). Apart from the slightly larger
grain size and the lower degree of abrasion, the zircon is identical in
all respects with the beach sand zircon.

River Sands. In the stream sands from the New England area,
almost perfect euhedral zircons, with intermediate diameters as great
as 8 mm., were handpicked from the concentrate. Those in the sands
from the granitic and sandstone areas are clear, and inclusions are not
abundant (Plate VI, figs. 1, 3 and 4).

Rutile.

Greywackes. The rare rutile grains are usually rounded, and smaller
than those in the beach sands. They are reddish-brown in colour.

Granitic Rocks. The range in colour from yellow-red to black
corresponds to that of the beach sand rutile. Most commonly the crystals
are deep reddish-brown. Although they are fractured during the
mechanical disintegration of the rocks, prism faces are often visible
indicating an elongate or stout habit, capped by pyramids.

Sandstones (Plate V, figs. 3 and 4). The rutile grains generally are
of a larger size and less abraded nature than those in the beach sands.
Some are euhedral or nearly so, and crystals edges usually can be seen.
Apparently owing to greater thickness, more grains are of a deeper
colour than those in the beach sands.

River Sands. Apart from the larger size and less abraded nature
of most of the grains (Plate VI, figs. 3 and 4), the rutile is identical
with that in the coastal sands.

Ilmenite.

Greywackes. Many of the grains, unlike those in the beach sands,
show partial alteration to leucoxene.

Granitic Rocks. In spite of fracturing from the mechanical dis-
integration, prism edges often can be seen.

Basalts. Grains fresh and show no alteration to leucoxene. The
lustre in reflected light is purple-grey and submetallic.

Sandstones. Usually larger and less abraded than those in the
beach sands. Some show alteration to leucoxene.

River Sand's. The grains commonly are ra,ther angular, particularly
in the samples from regions where Tertiary basalts are abundant
(Plate VI, fig. 2). Very few show partial alteration to leucoxene.

Garnet.

Greywackes. The rare grains of this mineral are smaller than from
beach sands. They are pink in colour and angular and irregular in shape.
Etching of the grain surfaces was not observed.

MINERAL BEACH SANDS OF SOUTHERN QUEENSLAND, PART II. 99

Granitic Rocks. The variety is almandinc, and the colour is pale
pink. Although fractured by mechanical disintegration, the dodecahedral
habit of the crystals is apparent. Inclusions are scarce.

Sandstones. Usually pale pink in colour, they are the same variety
(almandine) as occurs in the beach sands. Most are angular and irregular
in shape (Plate V, figs. 3 and 4), like those in the beach sands, but a
larger number show crystal edges and evidence of a dodecahedral habit
is sometimes seen. In etching and inclusions, they are identical with
the beach sand grains of this mineral.

River Sands. A dodecahedral habit is apparent in some, and crystal
edges are more often seen than in the beach sand garnet. Etching is
seen mainly on those from the sandstones areas (Plate VI, figs, 3 nnd 4).
They are commonly slightly larger than those in the beach sands, but
are similar in variety, colour and physical properties.

Tourmaline.

Greywackes. The grains are usually rounded, and some have a high
degree of sphericity. Prismatic crystals with the ends only slightly
rounded are also present. The most common colour is brown. Although
generally larger than the greywacke zircon, the size is smaller than the
beach sand tourmaline (Plate V, fig. 1).

Granitic Rocks. The tourmaline usually is brown, although grey,
blpe and green crystal fragments also occur, sometimes in the same
sample. Crystal prism edges are apparent.

Sandstones. The grains are not as rounded as those in the coastal
sands, and the short prismatic habit of the crystals is often seen
(Plate V, fig. 4). Colour and properties are identical with the beach
sand tourmaline.

River Sands. The grains are identical with those in beach sands.

Monazite.

Granitic Rocks. The monazite is honey-yellow in colour, and the
stumpy habit of the crystals is apparent. No mineral inclusions were
observed, but a few gas-filled cavities and minute dust-like inclusions
were seen.

Sandstones. Although most grains are rounded, a greater number
show crystal faces and edges than in the beach sands. They are honey-
yellow in colour, and often of fairly large size.

River Sands. The monazite is similar to that in the beach
sands, apart from larger grain size and less abraded nature (Plate VI,
figs. 3 and 4).

Leucoxene.

Greywackes. The grains are dull white in reflected light, and often
smaller in size than those in the beach sands.

Granitic Rocks. The very scarce leucoxene may have been derived
from the alteration of sphene which is common in sample G, the only
sample in which this mineral was found.

Sandstones and River Sands. The grains are similar to those in
the beach sands, apart from their slightly larger size and less rounded
nature.

100 PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND

Cassiterite.

Greywackes. The rare grains of this mineral are of very small size
and are not much rounded. They are particoloured reddish-brown
and colourless.

Granitic Rocks. A short bipyramidal tetragonal habit with striations
on the crystal faces is apparent. Some are zoned ; colours are ruby
and pale grey.

Sandstones and River Sands. The grains are similar to those in the
beach sands except for slightly larger size and less abraded nature.

Epidote.

Greywackes. The scarce grains are yellowish-green in colour, and
in various stages of alteration. They are faintly pleochroic.

Granitic Rocks. The prismatic habit usually is visible. Some grains
are clouded from alteration.

Sandstones and River Sands. The grains are often subhedral and
larger than those in the beach sands.

Magnetite.

Granitic Rocks. An octahedral habit is apparent in some, and the
material exhibits a silver-grey lustre in reflected light.

Sandstones and River Sands. Octahedral grains are sometimes seen.

Spinel.

Greywackes. The grains are small, green in colour and usually
subhedral.

Sandstones and River Sands. Like those in the beach sands, the
grains range from bright green to dark greenish-black in colour.
Apparently many are chrome spinel. They are slightly larger than
those in the beach sands ; some show octahedral habit.

Augite.

Basalts. The augite is usually the violet-tinted titaniferous variety,
but some grains are colourless or faintly green. Prism faces are usually
visible and inclusions of iron-ore are not uncommon.

River Sands. The grains are very small, and are prismatic with
rounded ends. Cleavage can be seen.

S phene.

Granitic Rocks. The sphene is brown in colour, and shows evidence
of a diamond or wedge-shaped habit. During the mechanical dis-
integration, fracturing has occurred, and irregularly shaped, ragged
particles have been produced. They have a vitreous lustre and some-
what watery appearance. Refractive index is particularly high, and
there is a total absence of extinction.

Sandstones and River Sands. The grains are usually fresher than
those in the beach sands, as well as being larger and less abraded.

Apatite.

Granitic Rocks. The prismatic crystals are colourless and clear to
brownish. Some show evidence of corrosion (probably magmatic).
Inclusions are rare, and so minute that they are indeterminable.

MINERAL BEACH SANDS OF SOUTHERN QUEENSLAND, PART II. 101

Sandstones and River Sands. The sparse grains are rounded and
very small.

Topaz.

Granitic Rocks. The topaz is colourless and glassy in appearance.
Fractured surfaces sometimes have a pale bluish tinge. Cleavage is
visible.

Sandstones and River Sands. The sparse grains are colourless and
rounded. They yield good biaxial, positive interference figures.

Anatase.

Greywackes. The grains are bluish, square and tabular, and show
no signs of wear. They are probably authigenic and derived in situ
from the decomposition of the ilmenite.

Granitic Rocks. The sparse anatase is yellow and indigo-blue in
colour, and the crystals have octahedral and square, tabular habits.
Some show zoning or geometric patterning. There is good cleavage.
Pleochroism is faint but distinct.

Sandstones. The occurrence of perfect crystals showing square,
tabular habit suggests that this mineral is authigenic.

Kyanite.

Greywackes. The grains are colourless. They are usually elongated
and show prism edges, with rounded ends. There is good cleavage.
Inclusions are rather common.

IX. CONCLUSIONS AND OUTLINE OF GEOLOGICAL HISTORY
OF THE HEAVY MINERALS.

For the first time, mechanical analyses of panned heavy mineral
concentrates and statistical measures derived from their cumulative
frequency curves have been used to compare and describe the physical
composition of beach sand concentrates. This has been possible because
the particular concentrates are made up almost entirely of minerals
which have similar specific gravities, and have been concentrated in
grains of very similar size.

Statistical measures, particularly the median diameter and Trask’s
coefficient of sorting, have been found to be very useful in the work
on the mechanical composition of the concentrates. By plotting the
median diameter and the coefficients of sorting and skewness against
the geographic distance along the coast, changes in these values can
clearly be seen, trends observed, and suggestions concerning their relation
to heavy mineral supply and direction of transport have been made
possible. From the abrupt increases in median diameter and coefficients
of sorting around the major coastline breaks and about the headlands
of Mesozoic sandstone, it is suggested that heavy mineral material has
been added to the shore at these points. This view is supported by
the decrease in the degree of mineral abrasion at these places.

The heavy minerals are well sorted but, contrary to common belief,
the highest degree of sorting is not present in the dune sand concentrates.

Over 90% by weight of the heavy minerals in all the coastal sand
samples is made up of zircon, rutile and ilmenite. Of the other minerals,
garnet, tourmaline, monazite and leucoxene are the more common. The
concentrates are distinguished from those in other parts of the world
by the high zircon and rutile content.

102 PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND

The most striking feature in distribution is a decrease in zircon and
increase in ilmenite content from south to north as far as the South
Passage. Northwards from the South Passage the percentages of zircon,
rutile and ilmenite remain more or less constant. The heavy mineral
analyses of the samples from localities south of the South Passage, in
which there is a higher zircon and rutile content, combined with the
results of the writer’s field survey, clearly indicate that the deposits in
that region are of greater economic value than those to the north.

For the enquiry into the source of the heavy minerals, study of
increases and decreases in content of the various minerals, presence
and absence of certain species, mineral appearance, grain size, and degree
of abrasion, have been found useful. It is concluded that the Mesozoic
sandstones of the region are the source of most of the heavy minerals
in the coastal sand deposits, but that at least some of the ilmenite has
been derived from the Tertiary basalts. The fact that the beach sand
ilmenite throughout the area usually is less abraded than the zircon and
rutile, in spite of its softer nature, supports the view that the detrital
history of this mineral generally has been shorter than that of the zircon
and rutile.

Although the sandstones are the immediate source rocks of most
of the heavy minerals, they are a secondary source. As they are of
Upper Triassic and Jurassic age, the primary source or sources of the
heavy minerals must be rocks of an earlier period. The heavy minerals
of the coastal sands suggest an igneous source, not metamorphic. Such
metamorphic minerals as kyanite and staurolite are almost absent from
the sands, although they are comparatively stable heavy minerals and
are abundant in beach deposits in other parts of the world. Many of
the species in the beach sands indicate an acid plutonic source. The
only rocks of this type of pre-Upper Triassic age in the region investi-
gated, as well as to the south, West and north, are the Permian
“ granites.” It is concluded that these granitic rocks are the primary
source of most of the heavy minerals. The pegmatites, greisens, and
other marginal products of the granitic bodies, including some rutile and
tin lode deposits, are believed to be of particular importance as source
rocks.

Following the uplift and earth movements at the close of the
Palaeozoic, the Permian granitic masses were soon unroofed, and with
weathering, there began the first cycle of erosion, transportation, and
deposition of most of the heavy minerals of the coastal sand deposits.
There was a large amount of erosion before the Jurassic. The marginal
parts of the granitic bodies would be first exposed to weathering, and
as these parts were rich in heavy accessory minerals, the swiftly flowing
streams draining the rejuvenated area would bring a large amount of
heavy minerals into the Mesozoic freshwater lakes. During this first
cycle of erosion, some of the less stable minerals would be removed, and
a greater concentration of more stable heavy minerals effected.

Since the draining of the lakes almost all the region has been dry
land. Much of the freshwater Mesozoic sandstones are soft and easily
eroded, and with weathering and disintegration, there began the second
cycle of erosion, transportation, and deposition of most of the heavy
minerals of the coastal sand deposits. The minerals were carried towards
the Pacific Ocean by coastal streams which were fast flowing, particularly
after the movements of uplift and consequent rejuvenation in Cainozoic
times. Also during this cycle a greater concentration of hard and

MINERAL BEACH SANDS OF SOUTHERN QUEENSLAND, PART II. 103

chemically resistant heavy minerals was effected by the removal of the
less stable species. The minerals generally became more abraded, and
the grain size finer.

Although there were outpourings of basic lava in Pliocene times
over large parts of the area, the main coastal streams are believed to
have existed in approximately their present positions before that time.
Following the major uplift at the close of the Tertiary, erosion was
particularly active and much detritus from the basalts was transported
seawards by the streams, and a much greater concentration of ilmenite
was effected than of the less stable augite and apatite. In addition to
the material brought down by the coastal streams, direct erosion of the
headlands of Mesozoic sandstone and Tertiary basalt along the coast has
supplied heavy detrital minerals to the shore. The higher ilmenite
content of the beach sand concentrates in the northern part of the region
is due to the greater amount of ilmenite brought down by the coastal
streams in that part. Most of this is believed to have come from the
Tertiary basalts, as the Mesozoic sandstones examined from the northern
areas do not have an ilmenite content appreciably greater than those
in the south.

During periods of coastal subsidence in Quaternary times the beach
and adjacent dune sands were passed through the surf zone. Also,
there must have been in the past, as at present, some direct erosion of
the sand deposits adjacent to the beach by high storm waves.

Although the heavy minerals are being brought down by coastal
streams at the present day, particularly during floods, most of the heavy
minerals in the coastal sand deposits reached the sea in pre-Recent times.
They have been moved northwards along the coast by beach and long-
shore drift, but they have not been carried very great distances.
Storm-wave and strong wind action have concentrated deposits, in some
cases of very considerable size and economic value.

X. ACKNOWLEDGMENTS.

This work was carried out at the Imperial College of Science and
Technology, London, during the tenure of an overseas studentship
awarded by the Commonwealth Council for Scientific and Industrial
Research.

I would like to thank Mr. J. G. C. Leech and Professor H. H. Read
of Imperial College, London, and Mr. R. G. C. Bathurst of the Sedgwick
Museum, Cambridge, for helpful discussion. I am also indebted to the
late Professor H. C. Richards, Professor W. H. Bryan and Dr. O. A.
Jones of the University of Queensland, under whose guidance the work
leading up to the present research was carried out.

I gratefully acknowledge the co-operation of the Queensland
Geological Survey in collecting and forwarding to me a number of beach
sand and rock samples, and of the Australian Bureau of Mineral Resources,.
Geology and Geophysics for a number of river sand samples. Thanks
are extended to Consolidated Zinc Corporation Ltd., for financial
assistance towards the cost of publication.

104

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND

XI.— BIBLIOGRAPHY.

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— (1905b).— “ The Geology of the New England Plateau, with special
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Boswell, P. G. H. (1933). — “ On the Mineralogy of Sedimentary Rocks.” Murby
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Bramlette, M. N. (1929). — “Natural Etching of Detrital Garnet.” Amer. Min.,
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Culey, A. G. (1932). — “ Notes on the Mineralogy of the Narrabeen Series of New
South Wales.” Proc. Roy. Soc. N.S. Wales, vol. 66, pp. 344-377.

Dryden, L., and Dryden, C. (1946). — “ Comparative Rates of Weathering of
some common Heavy Minerals.” J. Sed. Pet., vol. 16, pp. 91-96.

Fisher, N. H. (1945). — “ Titanium : Rutile and Ilmenite.” Aust. Min Res. Surv.
Sum. Rept. No. 2, pp. 1-21.

Krumbein, W C., and Pettijohn, F. J. (1938). — “ Manual of Sedimentary
Petrography.” Appleton-Century Co., New York.

Mackie, W. (1923). — “ The Principles that regulate the Distribution of Particles
of Heavy Minerals in Sedimentary Rocks, as illustrated by the Sandstones
of North-east Scotland.” Trans. Edin. Geol. Soc., vol. 11, Pt. 2,
pp. 138-164.

Miller, R. (1945). — “ The Heavy Minerals of Florida Beach and Dune Sands ”
Amer. Min., vol. 30, pp. 65-75.

Milner, H. B. (1929). — -“ Sedimentary Petrography.” 2nd Edition. Murby &
Co., London.

(1940). “ Sedimentary Petrography.” 3rd Edition. Murby & Co.,

London.

Morton, C. C. (1946).- — Verbal Communication, 22/2/46.

Pettijohn, F. J. (1941). — “ Persistence of Heavy Minerals and Geological Age.”
J. Geol., vol. 49, pp. 610-625.

Richards, H. C. (1916). — “ The Volcanic Rocks of South-Eastern Queensland.”
Proc. Roy. Soc. Queensl., vol. 27, pp. 105-204.

Trask, P. D. (1932). — “ Origin and Environment of Source Sediments of Petroleum.”
Gulf Pub. Co., Houston, U.S.A.

Twenhofel, W. H.,'and Tyler, S. A. (1941). — “ Methods of Study of Sediments.”
McGraw Hill Co., New York.

Tyler, S. A., and Marsden, R. W. (1938).—“ The Nature of Leucoxene.” J. Sed..
Pet., vol. 8, pp. 55-58.

Proc. Roy. Soc. Q’land., Vol. LXI., No. 7

Plate 1

Plate I.-^-Heavy Mineral Beach Sand ; Sample No. 7 (Tugun) x 35.

Fig. 1 : Bromoform-separated heavies. The rutile and iron-ore minerals appear
black in the photograph. Fig. 2 : Garnet fraction. Note large size, angularity,
and etched surface of grains. Fig. 3 : B.S.S. + 120 size fraction, after extraction
of highly magnetic and moderately magnetic minerals. Fig. 4 : B.S.S. — 120 size
fraction, after extraction of highly magnetic and moderately magnetic minerals.

Plate II. — Bromoform-separated Heavy Mineral Sands x 35.

Fig. 1 : Beach on North Stradbroke Island ; sample No. 21. Note two large,
translucent grains of tourmaline. Fig. 2 : Dunes on North Stradbroke Island ;
sample No. 27. Note great number of black grains ; also tourmaline grain showing
one prism-edge near lower left-hand corner. Fig. 3 : Beach at Caloundra Head ;
sample No. 39. Note large size of grains, especially of garnet, In this concentrate
from near sandstone headland. Fig. 4 : Beach at Point Arkwright : sample No. 41.
Same comments as for Fig. 3 apply. Note three grains of garnet towards left, and
the large, subhedral grain of monazite at right of centre.

Proc. Roy. Soc. Q'land., Vol. LXI., No. 7

Plate 3

Plate III. -Mineral Sand Concentrates from Tugun x 35.

Fig. 1 : Zircon concentrate, electrostatically separated. Note comparative
scarcity of inclusions. Fig: 2 : Rutile concentrate, electrostatically separated.
Note remnants of crystal edges on some grains. This photograph was taken with
a Process Pan plate, hence the lighter-coloured rutile grains appear translucent.
Fig. 3 : Monazite concentrate, separated by gravity and electromagnetic means.
Fig. 4 : Cassiterite sand concentrate, separated by gravity.

Proc. Roy. Soc. Q’Land., Vol. LXI., No. 7

Plate 4

Plate IV.

Fig. 1 : Burleigh Head, S.E. Queensland ; made up of Lower Palaeozoic
greywacke overlain by Tertiary basalt. Fig. 2 : Upper Triassic Bundamba sand-
stone outcropping at Caloundra Head. Fig. 3 : Looking south from Paradise
Caves (Noosa Head) along Coolum Beach. Point Arkwright in the distance.
Fig. 4 : Freshwater Jurassic sandstone, showing current bedding, at Noosa Head.

Proc. Roy. Soc. Q’land., Vol. LXI., No. 7

Plate 5

Plate V. — Bromoform-separated Heavy Mineral Fractions x 35.

Fig. 1 : From Burleigh Head greywacke ; sample A. Note small size of grains,
cloudy appearance of zircon, and prismatic habit of some of the tourmaline grains.
Fig. 2 : Non-magnetic heavies from Tenterfield granite ; sample D. Note high
zircon content, and prismatic apatite and rutile fragments (black in photograph).
Fig. 3 : From Clarence Series sandstone ; sample N. Note the larger size of most
of the grains, their less abraded nature, and the poorer degree of sorting, as com-
pared with most of the beach sand concentrates. The long, prismatic grain near
the left-hand edge is rutile. Fig. 4 : From Jurassic sandstone at Point Arkwright ;
sample T. Note similarity in grain size with Point Arkwright bea.ch sand con-
centrate (Plate II, fig. 4), and lower degree of abrasion and sorting. A subhedral
grain of tourmaline is in lower part of photograph,

Plate 6

Proc. Roy. Soc. Q’land., Vol. LXI., No. 7

Plate VI. — Heavy Mineral Fractions from River Sands x 35.

Fig. 1 : In New England granitic area ; sample W. Note high content of
zircon and iron-ore minerals, and small degree of abrasion. Compare size of zircon
with Plate III, fig. 1. Fig. 2 : From Currumbin Creek; sample ZC. Almost all
the black grains are ilmenite. Note very low degree of abrasion. Fig. 3 : From
Oraro River, which flows entirely through Clarence Series sediments; sample U.
Note larger size, and less abraded and less well-sorted nature of the grains than
in most of the beach sands. A large, bean-shaped grain of monazit.e is at the
lower margin of photograph. Fig. 4 : From tributary of Richmond River ; sample Z.
As in Fig. 3 note large size of monazite and other mineral grains, and greater number
of euhedral and subhedral zircon grains.

FREDERICK MANSON BAILEY

Vol. LXI., No. 8.

105

MEMORIAL ADDRESS :

F. M. BAILEY : HIS LIFE AND WORK

By C. T. White, Government Botanist.

( Delivered before the Royal Society of Queensland, 28 th November, 1949 ;
issued separately ) .

Frederick Manson Bailey, the most outstanding member of a line
of botanists and horticulturists, was born at Hackney, England, on the
8th March, 1827, and died at Brisbane on the 25th June, 1915. His
father, John Bailey, had been with Conrad Loddiges and Sons, the
famous, nurserymen of Hackney, England, in the early part of the
nineteenth century. At that time, great interest was taken by British
gardeners in the new and intriguing flora of South Africa, on the one
hand, and Australia on the other. South Africa gave to the horticultural
world the “ Cape-bulbs ” which included numerous species of Gladiolus,
Waisonia, Ixia, Sparaxis and others which have been hybridised and
remained firm favourites in all parts of the world where they can be
grown. The South African heaths were another group introduced into
general culture about that time.

Australian plants for the most part require the protection of a coel
or temperate house in Britain, but became firm favourites, especially the
Acacias or wattles, Boronias, Leptospermums and other Myrtaceae and
the shrubby Proteaceae. The first half of the nineteenth century was
an active era in the publication of works with coloured plates of plants
and those of New Holland, as Australia was then most generally known,
claimed considerable attention.

In 1787, Wm. Curtis started the Botanical Magazine, a work still
regularly appearing and edited at the Royal Botanic Gardens, Kew
(Eng.), for the Royal Horticultural Society. The early volumes, like
those of the present day, contain numerous illustrations of Australian
plants. A few years later (1791) the first volume of H. C. Andrew’s
Botanist’s Repository appeared ; this work ran to 10 volumes, the last
appearing in 1849. Another important series was the Botanical Register,
commenced in 1815 and discontinued in 1847. It contained, especially
from Vol. XV onwards, when John Lindley assumed the editorship,
numerous coloured illustrations of Australian plants, particularly those
of the “Swan River Colony” (Western Australia). A more popular
series stressing methods of cultivation was Paxton’s Magazine of Botany ,
16 volumes, 1836-1849.

Two important works with coloured illustrations were J. E. Smith’s
A Specimen of the Botany of New Holland, published as early as 1793,
and R. Sweet’s Flora Australasica in 1827-28. In France in the years
1803-4 appeared Ventenat’s Jardin de la Malmaison, with very large
coloured plates, in which some Australian plants were described and
figured for the first time.

Conrad Loddiges and Sons was a firm of botanical nurserymen who
helped to introduce and popularise plants from abroad, especially South
Africa and Australia. In addition to being expert nurserymen, the
Loddiges were good botanists and published a notable work, The Botanical
Cabinet. This work, which is still regularly quoted in botanical literature.

106 PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND

consists of 20 volumes ; the first appeared in 1817, the last in 1833.
Each volume consists of 100 plates partly in colour ; thus 2,000 exotic
plants were figured, many for the first time. It was natural therefore
that a member of such a firm should be chosen by Governor Gawler as
Government Botanist and Curator of a Botanic Garden which it was
proposed to establish in the young colony of South Australia. For this,
John Bailey was to receive the magnificent salary of £80 per annum
With his family — wife, two sons, one of whom was Frederick Manson,
and I think a daughter— he left England on the 12th December, 1838,
in the frigate “Buckinghamshire” which cast anchor in Holdfast Bay,
South Australia, just 100 days later, 22nd March, 1839. She was the
largest vessel to visit Australia up to that time, her gross burthen being
1500 tons, and she made an exceptionally fast passage.

A botanic gardens was laid out on the banks of the Torrens River.
It was divided into two sections, one devoted to flower-beds and
ornamental plants, the other to an economic section. One important
function of the latter was the growing of fresh vegetables to combat
scurvy. I do not know when this Botanic Gardens was abandoned, but
it was probably towards the end of 1841 when drastic cuts in administra-
tion costs were made by Grey in an endeavour to pull the colony out of
bankruptcy. No money being available to carry on the Botanic Gardens,
John Bailey turned his hand to farming and later to nursery work,
establishing the Hackney Nursery, after which the present suburb of
Adelaide is named. In this venture his two sons were associated with
him under the name of John Bailey & Sons.

The early Botanic Gardens had no connection with the present one
in Adelaide, which was not established until 1855. F. M. Bailey was
somewhat annoyed, that in any subsequent history of South Australia,
1855 was given as the date of the foundation of the Botanic Gardens
and the earlier one ignored. The early Gardens, however, from what I
gathered from him in conversation, were largely devoted to the growing
of flowers, vegetables and field crops. An important part of the work
was the introduction of economic plants. One of the most important
of these was the true Zante currant. Life in the colony was hard in
the early forties. Wheaten flour was very expensive and Mrs. Bailey
used the grain of one of the Sorghums for the making of johnny cakes
and dampers. The grain was ground in the family coffee-grinder. The
skinless barley, according to the younger son, F. Manson Bailey, in one
of his reminiscences, was experimented with but had to be given up,
not only for fear of damaging the mill, but because the noise startled
the natives in their wurlies ! Lighting oil also was scarce, and frequently
the younger members were sent out to collect large heaps of small bark
strips with which they constantly fed the open fire so that the older
ones could read by the light given out by the blaze. Young Fred Bailey
had just turned 12 when the family arrived in South Australia, so that
it is only natural that — apart from an early schooling at the Foundation
School of the Independent Church at Hackney, England — he was largely
a self-taught man. He was an omnivorous reader and part of his reading
was done when ploughing ; this was made possible by the fact that
slow-moving bullocks instead of horses were used for this work. Partly
influenced by family reasons, and partly by the lure of the goldfields
being opened up in the fifties, F. M. Bailey (now a young man in the
twenties) decided to dissolve the partnership with his father and brother
and engage in mining. He had hardly time to try his luck, however,

F. M. BAILEY : HIS LIFE AND WORK

107

before he was recalled to Adelaide by the illness of his father. He
resumed his position in the business, but in 1858 resolved to go forth
on a fresh quest to New Zealand, where he took up land in the Hutt
Valley. Soon afterwards he decided to return to Australia. After a
brief stay in Sydney, he came to Brisbane in 1861 and was destined to
play a most important part in the scientific life of the new State where,
except for brief visits to the other States, New Zealand, and New Guinea,,
he was not to leave again during his long life.

Soon after landing in Brisbane he opened a seed store in Edward
Street, and in addition he collected botanical specimens for sale to British
and foreign Botanical Museums and Herbaria. Partly owing to times of
financial stress in the colony, and probably to a lack of business acumen,
the store was closed and he had to look elsewhere for a means of support
for himself and his family, by this time totalling six children | (three
sets of twins).

Official Life.

In 1875 the Government set up a board to enquire into the causes
of the disease of livestock and plants in Queensland and F. M. Bailey
was appointed botanist. On this cause he travelled extensively in the
State, especially in connection with plants reputed poisonous to livestock
on the one hand and with grasses and native pasture herbage on the
other.

In December, 1880, on the recommendation of his very close friend,,
the Rev. J. E. Tenison-Woods,, S.J., he was appointed Acting Curator
of the Queensland Museum, a position he held until March, 1882, when
a permanent Curator (Mr. C. W. De Vis) was appointed. The Report
of the Trustees of the Queensland Museum for the year 1881-82 contains
the following reference: — •

“ From December, 1880, until March, 1882, Mr. F. M. Bailey performed the
duties of Curator, carrying out the general management of the Museum together
with his own botanical work, in a manner highly satisfactory to the Trustees ;
and, by his efforts, the phytological specimens in the upper story had been put
in order.

During the last year the extensive botanical library, previously kept in the
Curator’s cottage at the Botanic Gardens, has been transferred to the large room
in the bagement floor of the Museum .building, where Mr. Bailey now works as.
Government Botanist. In this room are also contained the few works of reference
belonging to the Museum and the varied library of the Philosophical Society.”

In 1881, while still Acting Curator of the Museum, he was appointed
Colonial Botanist, a position he held until his death. He retained
quarters at the Queensland Museum until 1889, when the Department
of Agriculture, under whose auspices he worked, was transferred to a
building of its own in William Street. He stayed there until 1912, the
Museum of Economic Botany and the Herbarium occupying three large
rooms, one of which is the present Ministerial office. In 1912 the Museum
and Herbarium were transferred to a special building in the Botanic
Gardens, which was then a Government activity under the control of
the Department of Agriculture and Stock. On the inauguration of the
Greater Brisbane Scheme in 1925 the control of the Botanic Gardens
was transferred to the Brisbane City Council.

Bailey never received a high salary ; for many years he received £300
per annum, and I do not think this was ever exceeded. It was intended
to retire him from the Public Service in 1902, but he said he could not
accept retirement and would go on working as far as he could without

108 PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND

pay, and that if the authorities wished to retire him they would have
to carry him out into the street. After some publicity had been given
to the matter in the press and his work for the State publicly praised,
his services were retained but only on half-pay, £150 per annum, and
later, a few years before his death, this was raised to £200 per annum.

Journeys and Collections.

He was an ardent collector, and specimens of his gathering are well
represented in the Queensland Herbarium, while earlier ones especially
are in various herbaria abroad. For some years after his arrival in 1861
he collected assiduously about Brisbane. He was especially interested
in cryptogams, although apart from the pteridophytes he did little or
no work on these groups himself, but he corresponded and sent specimens
to various specialists as follows : — -

Fresh-water algae : — E. Askenasy, W. Schmidle and later M. Moebuis
and Otto Nordstedt. (Three of the Botany Bulletins (Nos. V,
VI and XI) issued by the Department of Agriculture, Brisbane,
dealt with these plants.)

Marine algae (seaweeds) These were not collected by Bailey in
his early days in Queensland, and it was not until a few years
before his death that he started sending them to A. D. Cotton
(Kew).

Characeae : — O. Nordstedt, and much later to H. & J. Groves
(London).

Fungi : — M. J. Berkley, later to M. C. Cooke, and then to George
Massee (Kew) ; in later years a few were determined by Miss
E. N. Wakefield (Kew). One of the earliest papers on Aus-
tralian Fungi was “List of Fungi from Brisbane, Queensland,
with Descriptions of New Species ,” by M. J. Berkley and C. E.
Browne. Based largely on F. M. Bailey’s collections, it was
read before the Linnean Society of London on 23rd March,
1878, and published in the following year in the Society’s
Transactions.

Lichens : — J. Mueller, later to Chas. Knight and J. Stirton.

Mosses V. J. Brotherus.

Liverworts : — F. Stephani.

Duplicates of most of these collections are in the Queensland
Herbarium though not all, because for some reason he always regarded
these as his private property and they were not incorporated in the
official collections until after his death. I am also afraid that the collec-
tions have been neglected for many years, largely due to the absence
in Queensland of specialists in most of these groups. Apart from local
botanising, his most extensive collecting trips as far as I can gather
were as follows : —

1873 : Rockingham Bay, Upper Herbert River, and Seaview Range,
North Queensland.

1876 : Warrego, Maranoa and Leichhardt Districts, Western Queens-
land. He apparently travelled to Roma, thence to Morven,
Augathella, Tambo, and on to Rockhampton.

1877 : Cairns and ranges about the Barron River, North Queensland.

F. M. BAILEY : HIS LIFE AND WORK

109

1886-1894 : The Field Naturalists’ section of this Society was formed
in 1886 and started with fortnightly excursions. From the beginning,
F. M. Bailey was one of the most active members of the section. The
years 1891-4 were particularly active botanically, as the members of the
section followed the Brisbane-Gympie railway line as it was being built
and collected extensively in the then rich rain-forests from Eudlo to
Cooroy. Many new species of plants were described by Bailey from this
region. The section, however, gradually dwindled until only three or
four members remained. The last report was one by F. M. Bailey on
some plants from Eumundi to Cooroy, read before this Society on
21st April, 1894, and published in Vol. X of its Proceedings (pp. 51-53).

1889 : Bellenden Ker Expedition (North Queensland). He always
regarded this as his main collecting trip. The results certainly added
considerably to a knowledge of the flora of tropical Australia. They
were published, including descriptions of new species, in the Report of
the Government Scientific Expedition to Bellenden Ker Range , which was
issued in two forms : (1) foolscap size as a Parliamentary Paper, and
(2) as a special publication in demy 8vo size of the Department of Agri-
culture and Stock, Brisbane. Both bear the date 1889 on the title page,
but it is probable that the first has priority. It is extremely rare,
however, even in Australian libraries, whereas the latter is moderately
common and was the one always quoted by Bailey himself and by
subsequent authorities. He republished several of the new species in
other publications, especially the third supplement to the Synopsis of
the Queensland Flora, and frequently gave this as the place of publication
with or without reference to the Report.

1895 : Georgina River, Western Queensland. This visit was made
especially to investigate the course of the so-called Georgina River disease
in cattle, thought to be due to a plant. He blamed Eremophila
hignoniiflora as the possible cause of the trouble. We now know this
to be one of the best of the inland fodder shrubs and .trees, and recent
investigations lay the blame on a member of the same genus, E. latrobei.

1897 : Torres Straits. Early in the following year, 7th January,
1898, he read a paper entitled A Few Notes on the Flora of the Islands
of Torres Straits and the Mainland about Somerset at the Sydney Meeting
of the Australasian Association for the Advancement of Science. This
was published in Vol. VII of the Association’s Proceedings.

1898 : British New Guinea. At that time the territory was governed
from Brisbane and F. M. Bailey accompanied Lord Lamington on a tour
of inspection. Facilities for collecting were poor and Bailey suffered
considerably from fever during the trip. The botanical results were in
consequence rather meagre, and the specimens collected for the most
part very poor. A complete list, without any descriptions of the plants
collected, was given under the title Names of easily Recognised Plants
observed by Lord Lamington s Party during New Guinea Expedition as an
appendix to a Parliamentary Paper, Report of Visit to New Guinea,
Brisbane, 1898. Descriptions of the new species appeared in three small
papers in the Queensland Agricultural Journal for September, October
and November, 1898.

In some ways he was not unlike his more famous contemporary
Ferdinand Mueller, in as much as he described many of his finds direct
in the field. As far as I could gather from conversation with him and
an examination of his field notebooks, he apparently sometimes sat

110 PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND

down on a log or in camp and described his specimens, at least those
he thought previously undescribed, in sufficient detail to use the held MS.
without alteration for his reports and publications. I should say it is
doubtful if many of these specimens were checked over with previous
collections, floras or monographs. Both he and Mueller had the habit
at times of naming new species from mere fragments ; for instance,
Mueller named Flindersia papuana from an immature fruit and Bassia
maclayana from a single seed, and Bailey named Myrtus metrosideros
from leaves only. Though both founded institutions that have become
centres of research in systematic botany, I doubt if either took a great
deal of interest in herbaria as such. Probably neither realised the
importance of type-collections or of full details on herbarium sheets, for
in both the National Herbarium at Melbourne and the Queensland
Herbarium, Brisbane, there are numerous specimens lacking collector’s
names, and dates and localities are often abbreviated. Both Mueller
and Bailey relied on their own knowledge of handwriting and general
“look” of a specimen to tell the collector’s name. Both had the habit
of drawing up descriptions of new species from composite collections,
but in this and other points mentioned they were to a large extent
characteristic of their time. Even in the largest herbaria of those
days, the collectors represented were comparatively few and curators
relied on their personal knowledge to associate the specimens in their
charge with collector and locality.

What I have read of Baron Mueller’s life and my personal contact
with F. M. Bailey lead me to the conclusion that both thought they
had “cleared up” the flora of Australia in general and Queensland in
particular respectively, leaving only a few crumbs for those who came
after. As a matter of fact, genus after genus of Australian plants is
literallyj“crying out” for revisional work in the light of modern knowledge,
and it will require many years and a host of workers before this work
is anything like complete.

Publications.

F. M. Bailey was a prolific writer, and from the date of his appoint-
ment as Colonial Botanist in 1881 until his death in 1915, scarcely a
day passed without some manuscript, however short, being prepared for
publication. He always had a particular liking for the ferns and their
allies, the lycopods, and before his official, appointment had published
privately as early as 1874 his Queensland Ferns, a book of 70 pages
and 63 small illustrations distributed over 23 plates at the end of the
work; these latter were made by his friend Mr. H. S. Eaton, a well
known local artist of the time.

While botanist to the Board set up by the Government to enquire
into the causes of diseases in live-stock in Queensland he published in
1878, in association with K. T. Staiger, then Government Analyst,
Vol. I of an Illustrated Monograph of the Grasses of Queensland ; 42 grasses
were described by Bailey and each species illustrated by electrotype
nature-printing by Staiger. This work is now very rare, and no sub-
sequent volumes were issued.

Before his appointment as Colonial Botanist he had prepared his
only work dealing with Australia as a whole. This was the Fern World
of Australia, published by Gordon and Gotch in 1881. It contains 106
pages and two plates, one illustrating the tribes of ferns as then under-
stood, the other the four genera of lycopods. It is not surprising that

F. M. BAILEY : HIS LIFE AND WORK

111

he dealt with the ferns of Australia as a whole, including Lord Howe
Island, for of the 245 species dealt with, only 30 are not found in Queens-
land although many are common to Queensland and the other States,
particularly northern New South Wales. He contributed several papers
of a general nature, such as some on Queensland fungi, medicinal plants
and introduced plants, to the early volumes of the Proceedings of the
Linnean Society of New South Wales. ,The most important was a Census
of the Flora of Brisbane (in conjunction with the Rev. J. E. Tenison-
Woods), read before this Society on 20th March, 1879, and published
in Vol. IV of its Proceedings. From the date of his appointment as
Colonial Botanist all his papers (with a very few exceptions in the
Proceedings of this and other Australian Societies) appeared in official
bulletins or journals of the Department of Agriculture (later Department
of Agriculture and Stock), or as special publications of that Department.
One exception was the Weeds and Suspected Poisonous Plants of Queens-
land, which appeared in 1905-6, and is specially mentioned below. A
list in chronological order (with notes) of his main works follows

1874 — Queensland Ferns.

1878 — An Illustrated Monograph of the Grasses of Queensland Vol. 1 (in con-
junction with K. T. Staiger). No other volumes were issued.

1881 — Fern World of Australia.

1883- — A Synopsis of the Flora of Queensland. Three supplements were sub-
sequently issued in 1886, 1888 and 1890 respectively. An appendix to
the synopsis that was also issued separately was A Classified Index
of the Indigenous and Naturalised Plants of Queensland. With the
additional supplements, fresh classified lists were issued as separate
publications and called Catalogue of the Indigenous and Naturalised
Plants of Queensland.

1885 — Catalogue of Plants in the Two Metropolitan Gardens , The Brisbane Botanic
Gardens and Bowen Park. This is more than a mere list as it gives
native countries and notes on properties. It is still a very useful guide
to the ornamental plants grown in Queensland.

1886, 1888, 1899 — Catalogue of Queensland Woods (Three editions). F. M.
Bailey was very active in getting together for display purposes collec-
tions of woods, grasses and economic plants for important exhibitions.
These were accompanied by descriptive catalogues. The most important
are those describing the timbers. Catalogues were compiled for
the following exhibitions : Colonial and Indian Exhibition, London,
1886 ; Centennial International Exhibition, Melbourne, 1888 ; and
Greater Britain Exhibition, London, 1899. The catalogues were issued
Separately in booklet form.

1887 —Plants Reputed Poisonous to Live Stock in Queensland (in conjunction
with P. R. Gordon). This is a book of 112 pages and 45 full-page
plates from excellent pencil drawings by Miss M. A. Hope.

1890— 1913 — Botany Bulletins (. Department of Agriculture, Brisbane), 1-17. The

first three bear only numbers of the general bulletin series of the Depart-
ment and are numbered Bulletins 4, 7 and 9 respectively. The only
indication that Nos. 7 and 9 are the second and third of the botanical
series respectively is in the prefatory notes. The next four were entitled
Bulletins 13, 18, 20 and 21, but in addition bore the sub-titles of Botany
Bulletin Nos. IV, V, VI and VII respectively. From then they were
simply entitled Botany Bulletin VIII and so on to XVII. The series
was continued by subsequent Government Botanists, but ceased with
No. XXII (1920).

1891 — Official Guide to the Museum of Economic Botany, Department of Agri-
culture, Brisbane.

1892 — Lithograms of the Ferns of Queensland. This work consists of 191 plates
of nature-prints direct from the stone ; in the case of some of the smaller
species more than one appears on a page. No letter-press accompanied
the plates.

112

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND

1893 — A Companion for the Queensland Student of Plant Life.

1894 — Botany Abridged.

1895 — A Half-Century of Notes for the Guidance of Amateur Fruit Growers ;
to which is appended A Comprehensive list of the Fruit-bearing Plants
from Time to Time Introduced into Queensland. Bull. No. 5 (2nd ser.).
Department of Agriculture, Brisbane.

1897 — A Companion for the Queensland Student of Plant Life and Botany
Abridged. A second edition of two works issued separately in 1893
and 1894 respectively.

1897-1915 — Contributions to the Flora of Queensland and Contributions to the
Flora of British New Guinea. A series of papers (mostly small) issued
in the Queensland Agricultural Journal. These Contributions were
commenced in the first issue of the Queensland Agricultural Journal
in July, 1897, and the last appeared posthumously in July, 1915.

1899-1902 — The Queensland Flora. This work consists of six volumes, the first
published, in 1899 and the last in 1902. Bailey had an admiration for
George Bentham, the author of the Flora Australiensis, that amounted
almost to worship. He did not alter Bentham’s descriptions except in
a very few cases, so the work is largely a compilation from the Flora
Australiensis plus additions made to the known flora of Queensland by
Mueller and Bailey himself. A General Index to the work was published
as a separate volume three years later (1905) from finance supplied by
a visitor to Australia, Mr. F. E. Klotten of Frankfurt-am-Main, Germany.

1907 — The Weeds and Suspected Poisonous Plants of Queensland. This was a
private venture and appeared in 15 monthly parts. In bound volumes
the title page is dated 1906, but the last part did not appear until the
following year.

1912 — Comprehensive Catalogue of Queensland Plants. This was to have been
simply a list like previous catalogues (see under 1883 — Synopsis of the
Queensland Flora), but the then Governor of Queensland, Sir Wm.
MacGregor, recommended to the Government the advisability of illustrat-
ing each family by at least one illustration. This was agreed to, so the
work is illustrated by 970 line figures mostly by the writer of the present
biographical sketch and 16 coloured plates by Mrs. Ellis Rowan. No
date appeared on the title page, the “ Prefatory and other Notes ” are
dated 1909, but a special one prefacing Addenda and Corrigenda is
dated 19th December, 1912. The first few copies were received from
the printer before Christmas of that year, though the general issue was
not made until early 1913. The work forms a useful illustrated
companion to The Queensland Flora. It included the cryptogams.

Association with Scientific Societies.

F. M. Bailey took a great interest in the scientific life of Australia
and was elected an Honorary Member of the Philosophical Society of
Queensland in January, 1884. He was a member of the first council of
this Society and later (1890-91) President. He was an Honorary
Corresponding Member of the Royal Societies of Victoria, Tasmania
and South Australia. In 1902 the Royal Society of New South Wales-
awarded him the Clarke Memorial Medal for outstanding researches in
Natural Science. He was elected a corresponding member of the Linnean
Society of New South Wales in 1879 and contributed several papers to
the earlier volumes of the Proceedings of that Society. He was a fairly
regular attendant at meetings of the Australasian (now Australian and
New Zealand) Association for the Advancement of Science, and was
president of Section D (Biology) at the thirteenth meeting held at Sydney
in January, 1911. He was elected a corresponding member of the
Pharmaceutical Society of Great Britain in 1892, of the Societe Royal
Botanique de Belgique in 1897, and of the Royal Botanical Society of
Edinburgh in 1905. He joined the Linnean Society of London in 1878
and retained his Fellowship till the day of his death.

F. M. BAILEY : HIS LIFE AND WORK

113

Personalia.

In appearance F. M. Bailey was a distinguished looking man, and
for at least the last forty years of his life wore a long beard but no
moustache. He was of rather sparse build and had drooping eyelids
which towards the end he had constantly to prop up. He always wore
soft shirts and a narrow black bow-tie. It is doubtful if he ever possessed
a dinner or evening suit in his life. I remember on one occasion an
invitation from a scientific society for a conversazione being received
with “Evening Dress” in small type at the bottom. He tore the invita-
tion into fragments in a rage and threw it into the waste-paper basket.
He was exceedingly frugal in habits and had a positive horror of
drunkenness, gluttony and bawdy conversation. I remember him going
to Government House when (I think) Lady MacGregor said : “Well,
Mr. Bailey, I have got your favourite lunch — a stale bun” ! He never
took a drink of cold water so long as I knew him, as he said it gave him
a headache. This must have been a life-time habit, for when collecting
in the bush, if facilities for boiling the billy and making tea were not
available, he dipped his lunch — probably a stale bun — in a nearby creek.
Every night for many years he treated himself to three teaspoonsful of
rum — no more or less — in a glass of water. He rarely — at least in later
life — took other forms of alcohol. He never lost interest in the practical
side of horticulture, and one of his earliest publications issued by the
Department of Agriculture was A Half Century of Notes for the Guidance
of the Amateur Fruit Grower. Like many of his contemporaries in Aus-
tralian science, he took little interest in the philosophical side of his
subject, and Darwinism and all that followed it passed him by, making
little or no impression. I suppose there was so much at hand of direct
interest, and Europe in those days must have seemed a long way away,,
that this attitude in Bailey and others is not to be wondered at. He
never spread himself to sciences other than botany — at least to any
extent — though when Acting Curator of the Queensland Museum he
identified birds and mammals for the public as far as the library and
collections allowed. He also collected beetles for his friend and distant
relative, the celebrated coleopterist, Rev. Thos. Blackburn. He acted
on the principle of his favourite poet, Alexander Pope, who wrote:
“One science only will one genius fit, so vast is art, so narrow human
wit.” He was a great reader of poetry, and for years never went to
bed without a book of poems under his pillow, though it was the biting
wit of Pope or the more subtle one of Goldsmith that appealed most to
him. He could quote both authors at length. Pope’s contemporary
and friend, John Gay, was also quoted in the same way. Although he
read them fairly assiduously, poets such as Wordsworth and Tennyson
made little real appeal to him. He was, however, a great reader and
admirer of Keats and Byron, but he did not quote them extensively.
He was a man of extraordinary strong personality with a most lovable
and kindly character which was sometimes imposed upon. Indicating
his character, it may be mentioned he was a great friend of a German
medical man and scientist here, and one time a prominent member of
this Society — Dr. Jos. Lauterer. The latter was a great character in
his day, and once took a trip round the world. He carried a photograph
of F. M, Bailey with him, hung it up in his cabin or room, bowed to it
every day, and said “Good morning, Mr. Bailey.” Another great friend
was Mr. J. H. Simmonds, one time Hon. Secretary of the Field Naturalists
Section of the Royal Society and a keen local botanist and conchologist.
For many years Mr. Simmonds visited the Bailey grave on the anniversary

114 PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND

of F. M. Bailey’s death and laid a wreath upon it. These instances
indicate the tikeableness of a great man.

In religious matters, he was brought up an Independent, but after
his marriage in September, 1856, to Anna Maria, the daughter of the
Rev. Thos. Waite, an Anglican Clergyman, he attended the Church of
England for many years. With age, he became rather unorthodox, rarely
attended religious services, and did not claim allegiance to any particular
Christian sect. He always dubbed himself a “Deist” and in his own
words, “Saw in all nature the work of an Almighty hand.” He. was
created a C.M.G. in 1911. He was buried in the South Brisbane Cemetery.
A single daughter, Julia, who kept house for him for many years, was
later buried in the same grave. Four of his children survived him, three
daughters and a son. The latter was Director of the Botanic Gardens,
Brisbane, for some years and for a short time held the dual position of
Director of the Botanic Gardens and Government Botanist. Later he
was Director of the Botanic Gardens, Adelaide, from which position he
retired in 1932. One of his sons, Frederick Manson, is Chief Sylviculturist
of the New South Wales Forestry Commission, and another, John Rayer,
Curator of the- Botanic Gardens, Brisbane. The present writer, who
holds the position of Government Botanist of Queensland, is a grandson
on the maternal side.

The Royal Society of Queensland
REPORT OF THE COUNCIL FOR 1948

To the Members of the Royal Society of Queensland.

Your Council has pleasure in submitting the Annual Report of the
Society for the year 1948.

At Ordinary Meetings throughout the year four addresses were
given and one symposium held ; one meeting was devoted to exhibits,
and one to the presentation of papers. The Annual Memorial Lecture,,
held this year in honour of Dr. Alfred Jefferis Turner, was delivered
by Dr. I. M. Mackerras.

Ten original papers were accepted for publication in the Proceedings.

As Members are aware, the Annual Subscription has been raised
to twenty-five shillings for Ordinary Members and twelve and sixpence
for Associate Members. It is uncertain whether this will cover the
increased printing costs, and the Council is considering other possible
means of increasing income.

The Council has devoted considerable time to arranging the Library.
This work is almost complete, but revision of the catalogue is still
necessary.

There are 5 honorary life members, 9 life members, 3 corresponding
members, 220 ordinary members, and 1 associate member in the Society.
This year the Society has lost 2 members by death and 4 by resignation ;
22 ordinary members and 1 associate member have been elected. The
Council decided to remove from the list of members 38 who were several
years in arrears with subscriptions (vide Rule 15).

Attendance at Council Meetings was as follows : — H. C. Webster, 9 ;
E. M. Shepherd, 7 ; D. Hill, 5 ; M. I. R. Scott, 10 ; E. N. Marks, 9 ;
B. Baird, 6 ; S. T. Blake, 9 ; M. F. Hickey, 6 ; O. A. Jones, 9 ; A. L.
Reimann, 9 ; G. Mack, 9 ; J. H. Simmonds, 8 ; R. F. Langdon, 10.

HUGH C. WEBSTER, President.
Margaret I. R. Scott, Hon. Secretary.

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ABSTRACT OF PROCEEDINGS

VII.

Abstract of Proceedings, 28th March, 1949.

The Annual General Meeting of the Society was held in the Geology
Department of the University on Monday, 28th March, 1949, with the
President (Professor H. C. Webster) in the chair. His Excellency the
Governor and about sixty members and friends were present. The
minutes of the previous Annual General Meeting were read and confirmed.
The Annual Report was adopted and the Balance Sheet received.
Mr. D. H. Kemp and Professor W. V! Macfarlane were nominated for
Ordinary Membership and Mr. J. Green for Associate Membership.

The following officers were elected for 1949 : —

President : Dr. D. Hill.

Vice-President : Professor M. F. Hickey.

Hon. Secretary : Miss M. I. R. Scott.

Hon. Treasurer : Miss D. F. Sandars.

Librarian : Miss B. Baird.

Editors : Mr. S. T. Blake, Mr. G. Mack.

Members of Council : Dr. O. A. Jones, Mr. E. M. Shepherd,
Dr. A. L. Reimann, Mr. J. H. Simmonds, Professor L. J. H.
Teakle.

Hon. Auditor : Mr. L. P. Herdsman.

The Presidential Address, entitled “ Energy and the Future of
Mankind,” was delivered by Professor H. C. Webster. A vote of thanks
was moved by Professor W. H. Bryan, seconded by Mr. R. F. Langdon,
and carried by acclamation.

Abstract of Proceedings, 3rd May, 1949.

The Ordinary Monthly Meeting of the Society was held on Tuesday,
3rd May, with the President (Dr. D. Hill) in the chair. The minutes
of the previous meeting were read and confirmed. Professor W. V.
Macfarlane and Mr. D. H. Kemp were elected to Ordinary Membership ;
and Mr. J. Green to Associate Membership. Mr. T. J. Brockington,
Mr. W. G. Maxwell, Mr. P. J. O’Sullivan, Mr. A. K. Sutherland, Mr.
G. C. Simmons, and Mr. P. H. Durie were proposed for Ordinary
Membership.

Mr. V. Grenning gave an address entitled “ Forestry in Queens-
land,” of which the following is an abstract : — “ The timber supply
position in each State in Australia was briefly dealt with, the situation
in Queensland being treated in some detail. The main types of timber
being milled in Queensland are hardwoods and hoop and bunya pine
representing about three-quarters of the total quantity of timber milled.
The other classes are cypress pine, cabinetwoods and miscellaneous
species which are of importance but relatively small compared to the
two main classes. Approximately two-thirds of the timber cut comes
from Crown lands. The virgin hoop pine stands are rapidly disappearing
and at the present rate of cut will only last for a few years. In order
to make provision for future supplies, the Department has of recent
years embarked on a vigorous programme of softwood planting with the
object of establishing 200,000 acres as soon as possible. To date 40,000
acres have been established and planting is proceeding at the rate of

VIII.

ABSTRACT OF PROCEEDINGS

about 4,000 acres per annum, but it is hoped to increase this. About
two-thirds of the hardwood now being milled in Queensland comes from
private lands, which are not being protected and managed for the per-
manent production of this timber. The Department is taking action to
provide supplies for the future by re-establishing young forests on cut-over
country by natural regeneration, i.e., from the regeneration by natural
seed fall from seed trees. The cypress pine forests are re-established by
similar methods and to date approximately 500,000 acres of hardwood
and cypress pine forests have been given improvement and regeneration
treatment. Considerable extension of this work is projected. Reference
was also made to the methods adopted for protecting forests from fire
and the investigation and research work into the problems of reforestation
and utilisation was briefly mentioned.”

Abstract of Proceedings, 23rd May, 1949.

The Ordinary Monthly Meeting of the Society was held in the
Geology Department of the University on Monday, 23rd May, with the
President (Dr. Dorothy Hill) in the chair. The meeting was held in
conjunction with the Queensland Naturalists’ Club. About fifty members
of these Societies and friends were present. The minutes of the previous
meeting were read and confirmed. The following were elected to Ordinary
Membership : — Mr. T. C. Brockington, Mr. W. G. Maxwell, Mr. P. J.
O’Sullivan, Mr. A. K. Sutherland, Mr. G. C. Simmons, and Mr. P. H.
Durie. The following were nominated for Ordinary Membership —
Mr. D. J. Belford, Mr. J. N. Casey, Mr. W. L. Hawthorne, Mr. Grahame
Tweedale, and Mr. John F. G. Wilkinson.

Dr. O. H. Selling gave an address entitled “ Upper Cretaceous and
Tertiary Plant Remains in Antarctica.”

Abstract, of Proceedings, 27th June, 1949.

The Ordinary Monthly Meeting of the Society was held in the
Geology Department of the University on Monday, 27th June, with the
President (Dr. Dorothy Hill) in the chair. About eighty members and
friends were present. The minutes of the previous meeting were con-
firmed. The following were elected to Ordinary Membership : — Mr. D. J.
Belford, Mr. J. N. Casey, Mr. W. L. Hawthorne, Mr. Grahame Tweedale,
and Mr. John F. G. Wilkinson. Mr. K. S. W. Campbell was nominated
for Ordinary Membership.

A film on atomic physics was shown.

Abstract of Proceedings, 25th July, 1949.

The Ordinary Monthly Meeting of the Society was held in the
Geology Department of the University on Monday, 25th July, with
the President (Dr. Dorothy Hill) in the chair. About twenty-five
members and friends were present. The minutes of the previous meeting
were confirmed. Mr. K. S. W. Campbell was elected to Ordinary Member-
ship. Mr. B. E. Anderson was nominated for Ordinary Membership.

The following were exhibited

“ Quartz crystals,” by Professor W. H. Bryan.

ABSTRACT OF PROCEEDINGS

IX.

“ Some local commensal animals,” by Miss D. Sandars.

" Some eucalypts from the Northern Territory,” by Mr. S. T.
Blake.

“ Photographs from the Atomic Energy Commission,” by
Professor H. C. Webster.

“ Rocks from islands of Torres Strait,” by Mr. C. Ogilvie.

“ Rocks from the Logan district,” by Professor F. W.
Whitehouse.

“ Carbonized fruits from the Victorian deep leads,” by Mr.
F. S. Colli ver.

“ Photographs of a trypanosome from a Platypus,” by Dr. I. M.
Mackerras. These photographs were sent from Hobart by
Dr. C. A. Duncan.

Abstract of Proceedings, 29th August, 1949.

The Ordinary Monthly Meeting of the Society was held in the
Geology Department of the University on Monday, 29th August, with
the President (Dr. D. Hill) in the chair. About fifty-five members and
friends were present. The minutes of the previous meeting were read
and confirmed. Mr. B. E. Anderson was elected to Ordinary Membership.
Mr. R. D. Malcolmson, Mr. W. B. Mather, Mr. J. M. Thomson, Professor
M. Shaw, and Professor J. H. Lavery were nominated for Ordinary
Membership.

Professor W. V. Macfarlane gave an address entitled “ Human
Reactions to Atomic Radiations : A Survey of 500 Years.” The histories
of the major natural exposures of man to radiations were related. First
the bergsticht of Schneeberg (Saxony) miners from 1410 to the present
day was related to uranium products which produced carcinoma of the
lung. At Joachimsthal in Czechoslovakia a similar high death rate from
cancer of the lung was identified in 1926. Probably 3,000 miners had
died from this radium-induced disease in five centuries. At least 200
doctors, technicians, patients and nurses had died from skin-cancer
engendered by X-rays since 1895. At Orange in New Jersey from
1917-24, 19 cases of radium poisoning causing sarcoma of bone, or aplastic
anaemia were recorded. Finally, in 1945, the atom bombs on Japan
exposed thousands to acute radiation-injury — depilation, vascular
damage, and aplastic anaemia, with death from intercurrent infection.

The mechanism of these processes was considered. Essentially all
damage was due to ionisation which, if intense enough, destroyed living
protoplasm. In smaller doses the alpha, beta, gamma rays, neutrons or
protons ionised water or solutes to produce reducing or oxidising sub-
stances which destroyed enzymes and proteins. Other ions forming in
the nucleo-protein of chromosomes caused gene-mutations which were
lethal at the subsequent division. Chromosomes also broke or divided
irregularly or became adherent to each other and prevented normal
development. All these mechanisms help to account for the damaging
or therapeutic effects of radiations.

The carcinogenetic effects from e.g., short-range alpha-particles or
neutrons are more difficult to account for. It seemed likely that slow
changes were set up in cells by ions, which might act like other carcinogens
(amidofluorene) which sensitised the cells. The cells then might divide
erratically as the result of any mild irritant such as bronchitis or dust.

ABSTRACT OF PROCEEDINGS

Abstract of Proceedings, 26th September, 1949.

The Ordinary Monthly Meeting of the Society was held in the
Geology Department of the University on Monday, 26th September,
with the President (Dr. Dorothy Hill) in the chair. About seventy-five
members and friends were present. The minutes of the previous meeting
were confirmed. The following were elected to Ordinary Membership : —
Mr. W. B. Mather, Mr. R. D. Malcolmson, Professor M. Shaw, Professor
J. H. Lavery, and Mr. J. M. Thomson.

Mr. R. F. Langdon presented a paper entitled “A New Ergot from
Queensland.”

Two addresses were given. Professor J. F. Bonner spoke on
“ Recent Adventures in Plant Physiology,” and Professor K. C. Hammer
spoke bn “ Photoperiodism.”

Abstract of Proceedings, 31st October, 1949.

The Ordinary Monthly Meeting of the Society was held in the
Geology Department of the University on Monday, 31st October, with
the President (Dr. D. Hill) in the chair. About thirty-five members
and friends were present. The minutes of the previous meeting were
confirmed. Professor W. Stephenson and Dr. W. A. McDougall were
nominated for Ordinary Membership.

Mr. J. S. Just gave the meeting details of the Science House project
as far as it has gone, and after stating the names of the various interested
bodies advised that a “ Ways and Means ” Committee had been formed
on which were himself as Chairman, Professor Wilkinson, Mr. G. F.
Cuppaidge, Professor Cummings, and Mr. R. F. G. Wilson. The Com-
mittee had set out as an objective a building of approximately 9,000
square feet in area and containing a library with store room in the
basement, a large and a small lecture room, committee rooms, offices
and amenities block at a then estimated cost of £35,000. After dealing
with interviews and deputations which had taken place between members
of the Government and the Committee, Mr. Just went on to state that
it was anticipated that bodies such as ourselves and others should make
a levy on members amounting to, say, 25/- per member over five years
as a contribution towards the capital cost of the building. As over
7,000 members were involved a substantial basic sum would be assured.
Other bodies may care to put in investment over and above that levied
sum. The levied sum would be treated as normal capital, whilst other
investments may or may not be treated as debentures at the wish of
the interested bodies. It was also expected that donations from interested
persons and bodies would make a substantial sum. He pointed out
that tentatively the “ Ways and Means ” Committee had suggested
that the management should be in the hands of a Board of Directors
nominated by those who contributed towards the cost of the building
on the basis of one nominee for each £1,000 contributed. Bodies con-
tributing less than £1,000 could join together to elect their own repre-
sentative, but that was a matter for final discussion. Mr. Just also
drew attention to the Lord Mayor’s suggestion that a cultural centre
should be created at the foot of Albert Street, and added that no more
worthy object than Science House could be put forward as forming the
first building to form part of a cultural block in that centre. In connec-
tion with the use of the building, it was pointed out that as well as

ABSTRACT OF PROCEEDINGS

XI.

rentals from lecture rooms it was suggested that the combined libraries
of the constituent bodies should be housed on the one floor under a
competent librarian and that, whilst retaining their entirety, would be
available to all constituent body members. This library would form the
nucleus of the most important scientific and technical library within the
Commonwealth. This library, in itself, makes the effort well worthy of
Government support and it was hoped that help would soon be forth-
coming. Mr. Just answered several questions.

The following papers were presented

" Contributions to the Geology of Brisbane. No. 1. Local Applica-
tions of the Standard Stratigraphical Nomenclature,” by W. H. Bryan
and O. A. Jones.

“ Marine Insects,” by I. M. Mackerras.

Abstract of Proceedings, 28th November, 1949.

A Special Meeting of the Society was held in the Geology Depart-
ment of the University on Monday, 28th November, with the President
(Dr. Dorothy Hill) in the chair. The business of the meeting was the
appointment of a Trustee to fill the vacancy caused. by the resignation
of Mr. J. B. Henderson. On the motion of Mr. S. T. Blake, seconded
by Mr. J. H. Simmonds, Dr. E. O. Marks was appointed.

The Ordinary Monthly Meeting of the Society was held in the .
Geology Department of the University on Monday, 28th November, with
the President (Dr. Dorothy Hill) in the chair. About forty members
and friends were present. The minutes of the previous meeting were
confirmed. Professor W. Stephenson and Dr. W. A. MacDougall were
elected to Ordinary Membership.

The following papers were read by title : —

“ Additions to the Flora of Arnhem Land,” by C. T. White.

“ Revision of Bregmaceros with Descriptions of Australian
Larval Forms,” by I. S. R. Munro.

“ Heavy Mineral Beach sands of Southern Queensland, Part II,”
by A. W. Beasley.

Mr. C. T. White delivered a memorial lecture entitled “ F. M.
Bailey : His life and work.”

Professor F. W. Whitehouse exhibited a series of cephalopod faunas
from various horizons in Western Queensland ranging from the bottom
to the middle of the Ordovician, having close affinities with the faunas
of corresponding ages from the Boreal Province of the Northern
Hemisphere.

Mr. W. G. H. Maxwell exhibited brachiopods collected by Mr. J. H.
Reid, from beds occurring to the south of Mount Morgan. They belong
to the new genus, Cyrtospirifer reidi, and indicate the age of these beds
to be upper Devonian. These are the most southern Upper Devonian
marine beds which have been found in Queensland.

XII

ABSTRACT OF PROCEEDINGS

CHANGES IN MEMBERSHIP

New Members.

Anderson, B. E. ... ... ... c/-

Belford, D. J., B.Sc. ...

Brockington, T. J.

Campbell, K. S. W., B.Sc

Casey, J. N., B.Sc.

Durie, P. H., B.Sc. ...

Hawthorn, W. L., B.Sc.

Kemp, D. H.

Lavery, Prof. J. H., M.E., B.Sc.
MacDougall, W. A., D.Sc.
Macfarlane, Prof. W. V., M.A., M.D.
Malcolmson, R. D., B.Sc.

Mather, W. B.

Maxwell, W. G. H

O’Sullivan, P. J., B.Agr.Sc. ...
Shaw, Prof. M., M.Eng., M.Mech.E.,
M.I.Mech.E., A.M.I.E. (Aust.)
Simmons, G. C., B.Sc.

Stephenson, Prof. W., Ph.D.
Sutherland, A. K., B.V.Sc., M.S.
Thomson, J. M.,

Tweedale, G., B.Sc., ...

Wilkinson, J. F. G., B.Sc.

Green, J.

Cardno & Davies, New Zealand
Chambers, Queen Street, Brisbane,
c/- Australian Petroleum Company, Port
Moresby.

McNaughton Street, Redcliffe.

Geology Department, University.

Bureau of Mineral Resources, Geology and
Geophysics, Canberra^

Veterinary Parasitology Lab., C.S.I.R.O.,
Yeerongpilly..

Geological Survey, Brisbane.

56 Heath Street, East Brisbane.
Engineering Dept., University, Brisbane.
Dept, of Agriculture and Stock, Brisbane.
Physiology Dept., University.

Physics Dept., University.

Zoology Dept., University.

Geology Dept., University.

Animal Health Station, Yeerongpilly.

Engineering Dept., University.

Animal Health Station, Yeerongpilly.
Zoology Dept., University, Brisbane.
Animal Health Station, Yeerongpilly.
Fisheries Research Station, Dunwich.
Geology Dept., University.

Geology Dept., University.

Associate Member.

Botany Dept., University.

Earnshaw, Dr. P. A.
Ellis, C.

Fisher, Dr. E.
Fraser, C. S.
Hamlyn-Harris, R.

Resignations.

Lamberton, J. A.
McConnell, Miss U.
Mackenzie, A. D.
Macpherson, Dr. R.

Marks, Dr. A.
Phillips, Dr. B. J
Price, Dr. T. A.
K. Webb. L. J.

Death.
Bick, E.

GUIDE FOR THE PREPARATION OF SYNOPSES

1. PURPOSE.

It is desirable that each paper be accompanied by a synopsis preferably
appearing at the beginning. This synopsis is not part of the paper ; it is intended
to convey briefly the content of the paper, to draw attention to all new information
and to the main conclusions. It should be factual.

2. STYLE OF WRITING.

The synopsis should be written concisely and in normal rather than abbreviated
English. It is preferable to use the third person. Where possible use standard
rather than proprietary terms, and avoid unnecessary contracting.

It should be presumed that the reader has some knowledge of the subject
but has not read the paper. The synopsis should therefore be intelligible in itself
without reference to the paper, for example it should not cite sections or illustra-
tions by their numerical references in the text.

3. CONTENT.

The title of the paper is usually read as part of the synopsis. The opening
sentence should be framed accordingly and repetition of the title avoided. If the
title is insufficiently comprehensive the opening should indicate the subjects covered.
Usually the beginning of a synopsis should state the objective of the investigation.

It is sometimes valuable to indicate the treatment of the subject by such
words as : brief, exhaustive, theoretical, etc.

The synopsis should indicate newly observed facts, conclusions of an experiment
or argument and, if possible, the essential parts of any new theory, treatment,
apparatus, technique, etc.

It should contain the names of any new compound, mineral, species, etc., and
any new numerical data, such as physical constants ; if this is not possible it should
draw attention to them. It is important to refer to new items and observations,
even though some are incidental to the main purpose of the paper ; such information
may otherwise be hidden though it is often very useful.

When giving experimental results the synopsis should indicate the methods
used ; for new methods the basic principle, range of operation and degree of accuracy
should be given.

4. DETAIL OF LAYOUT.

It is impossible to recommend a standard length for a synopsis. It should,
however, be concise and should not normally exceed 100 words.

If it is necessary to refer to earlier work in the summary, the reference should
always be given in the same manner as in the text. Otherwise references should
be left out.

When a synopsis is completed, the author is urged to revise it carefully,
removing redundant words, clarifying obscurities and rectifying errors in copying
from the paper. Particular attention should be paid by him to scientific and
proper names, numerical data and chemical and mathematical formulae.

CONTENTS

Vol. LXI.

No. 1 — Presidential Address : Energy and the Future of Mankind.

By H. C. Webster, D.Sc ., Ph.D., F.Inst.P. (Issued separately,
30th December, 1950)

No. 3 — Marine Insects. By I. M. Mackerras, F.R.A.C.P. (Issued
separately, 30th December, 1950) ...

No. 4 — A New Ergot from Queensland. By R. F. N. Langdon,

M.Agr.Sc. (Issued separately, 30th December, 1950)

No. 5 — Revision of Bregmaceros with Descriptions of Larval Stages
from Australasia. By Ian S. R. Munro, M.Sc. (Issued
separately, 30th December, 1950)

No. 6 — Additions to the Flora of Arnheim Land. By C. T. White.

(Issued separately, 30th December, 1950) ...

No. 7— Heavy Mineral Beach Sands of Southern Queensland.

Part II. — Physical and Mineralogical Composition,
Mineral Descriptions, and Origin of the Heavy Minerals.
By A. W. Beasley, Ph.D., D.I.C., F.G.S. (Issued separately,
30th December, 1950) ... ...

No. 8 — F. M. Bailey : His Life and Work. By C. T. White. (Issued
separately 30th December, 1 950) ... ... ... . . <.

Report of Council

Abstract of Proceedings

Changes in Membership ... ...

Pages

1-11

13-18

19-29

31-35

37-54

55-58

59-104

105-114

v.-vi.

vii.-xi.

xii.

PROCEEDINGS

OF THE

ROYAL SOCIETY

QUEENSLAND

FOR 1950

VOL. LXII.

This volume includes the
C. T. WHITE MEMORIAL SUPPLEMENT

ISSUED 15th SEPTEMBER, 1952.

PRICE: TWENTY-FIVE SHILLINGS,

Printed for the Society
by

A. H. TUCKER, Government Printer, Brisbane.

The Royal Society of Queensland.

Patron :

HIS EXCELLENCY LIEUT.-GENERAL SIR JOHN D. LAVARACK, C.B.,
C.M.G., D.S.O., C. de G., K.B.E.

OFFICERS, 1950

President :

Professor M. F. HICKEY, M.A., M.B., B.S.

Vice-Presidents :

DOROTHY HILL, D.Sc., Ph.D.
Professor H. J. G. HINES, B.Sc.

Hon. Treasurer : Hon. Secretary:

DOROTHEA F. SANDARS, M.Sc. MARGARET I. R. SCOTT, M.Se.

Hon. Librarian : Hon. Editors:

F. S. COLLIYER. S. T. BLAKE, M.Sc.

G. MACK, B.Sc.

Members of Council:

I. M. MACKERRAS, D.Sc., Professor A. L. REIMANN, D.Sc., Ph.D.,
J. H. SIMMONDS, M.Sc., M.B.E., Professor L. J. H. TEAKLE,
Professor H. C. WEBSTER, D.Sc., Ph.D., F.I.P., F.R.M.S.

T r ustees :

F. BENNET, B.Sc., Professor W. H. BRYAN, M.C., D.Sc., and
E. O. MARKS, M.D., B.A., B.E.

Hon. Auditor:

L. P. HERDSMAN.

Bankers :

COMMONWEALTH BANK OF AUSTRALIA.

CONTENTS

Vol. LXII.

No. 1. — The Ordovician- Corals. By Dorothy Sill, D.Sc., Ph.D. (Issued
separately, 15th October, 1951) . . . . . -

No. 2. — Apistomyia collini Bezzi (Diptera, Blepharoceridae) in
North Queensland. By I. M. Mackerras and M. J.
Mackerras. (Issued separately, 5th November, 1951)

Technical Notes — Bandicoot Food. By Dorothea F. Bandars

C. T. WHITE MEMORIAL SUPPLEMENT.

No. 3. — Cyril Tenison White, 1890-1950. (Issued separately,
15th August, 1952)

No. 4. — Reductions in Elaeocarpus. By E. D. Merrill. (Issued
separately, 15th August, 1952)

No. 5.- — Vegetative Habit in the Genus Eulophia (Orchidaceae).

By B. E. Holttum. (Issued separately, 15th August, 1952)

No. 6. — Rheophytes. By C. G. G. J. van Steenis. (Issued separately,
15th August, 1952) . .

No. 7. — Pseudoraphis spinescens (R.Br.) n. comb., and some
Records of New South Wales Grasses. By Joyce TV.
Vickery. (Issued sejiarately, 15tli August, 1952)

No. 8. — The Significance of the Mallee Habit in Eucalyptus. By
N. T. Burbidge. (Issued separately, 29th August, 1952)

No. 9. — Opisthiolepis, a New Genus of Proteaceae from Queens-
land. By L. S. Smith. (Issued separately, 29th August
1952)

No. 10. — The Identification and Distribution of some Cyperaceae
and Gramineae, Chiefly from Australia. By S. T. Blake.
(Issued separately, 29th August, 1952)

No. 11. — Notes on Some Australian Compositae. By J. H. Willis.
(Issued separately, 22nd August, 1952)

No. 12. — Whiteochloa, a New Genus of Grasses from the Northern
Territory of Australia. By C. E. Subbard. (Issued
separately, 22nd August, 1952)

Report of Council

Abstract of Proceedings

Pages.

1-28

29-32

3£>-48

49-56

57-60

61-68

69-72

73-78

79-82

83-100

101-108

109-112

v-vi

vii-xiii

Changes in Membership

xiv

PROCEEDINGS

OF THE

ROYAL SOCIETY

QUEENSLAND

FOR 1950

VOL. LXII.

This volume includes the
C. T. WHITE MEMORIAL SUPPLEMENT

ISSUED 15th SEPTEMBER, 1952.

PRICE: TWENTY-FIVE SHILLINGS.

Printed for the Society
by

A. H. TUCKER, Government Printer, Brisbane.

NOTICE TO AUTHORS

1. Each paper should he accompanied by the author ’s name, degrees and official
address.

2. Papers must be complete and in a form suitable for publication when com-
municated to the Society and should be as concise as possible.

3. Papers must be accompanied by a synopsis prepared according to the direction
given on the inside of the back cover.

4. Papers should be in double-spaced typescript on one side of the paper with
ample margins.

5. The use of italics in the text should be restricted to generic and specific names,
foreign words, and titles of periodicals.

6. The cost of author’s corrections to proof above what the Council considers a
reasonable amount, must be borne by the author.

7. Unless otherwise specified each author will be supplied with fifty separate
copies of his paper. Any number exceeding this may be obtained at approxi-
mately cost price.

S. All references should be listed at the end of each paper and arranged
alphabetically under authors’ names, e.g.,

Keilin, D. (1929) Proc. Eoy. Soc. B., vol. 104, p. 207.

Lesage, P. (1895) Ann.Sci. Nat. Bot., vol. 1, p. 309.

The corresponding references in the text should be:

“ Keilin (1929) ”, “Lesage (1895) ”.

9. The size of the printed plate will not exceed 8 in. x 4| in., and drawings may
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in mind. Text figures should be drawn for reduction to a width not
exceeding 4 in.

Vol. LXII., No. 1.

Proceedings of the Royal Society of

PRESIDENTIAL ADDRESS.

By Dorothy Hill, D.Sc., Ph.D., Geology Department, University of

Queensland.

(Delivered before the Royal Society of Queensland, 21th March, 1950;
issued separately 15 th October, 1951.)

During the year, the Society lost a valued member by the death of
Mr. E. W. Bick.

Ernest William Bick was born in England, and came to Queens-
land with his parents as a child in 1884. He was employed by the
Queensland Acclimatization Society at its gardens at Bowen Park,
where he became propagator. He became horticulturist at Govern-
ment House in 1907, and later transferred to the Botanic Gardens
as collector. He went on his first collecting trip in 1910 to Torres
Strait and the Gulf of Carpentaria with Sir William MacGregor, and
later he collected in many parts of Queensland. On the retirement of
J. F. Bailey in 1915, he became Curator of the Brisbane Botanic
Gardens, a position which he retained until his retirement in 1940.

His membership of the Royal Society of Queensland began in
1918; he became Hon. Treasurer in 1922, and continued in that office
until 1947, when he was elected an Honorary Life Member in recogni-
tion of his outstanding service to the Society. He was for many years
President of the Horticultural Society of Queensland; he was a Past
President of the Queensland Orchid Society and the Queensland
Naturalists 7 Club, and a Vice-President of the Queensland Acclima-
tization Society at the time of his death. He also served as Hon.
Auditor of the Naturalists ’ Club, and of the Entomological Society of
Queensland. He represented Queensland as a nominee of the Depart-
ment of Agriculture and Stock at several meetings of the Australian
and New Zealand Association for the Advancement of Science. He
died in 1949, and is survived by his wife, son, and daughter.

THE ORDOVICIAN CORALS.

(With Text Figs. 1-4.)

I. INTRODUCTION.

The Ordovician corals may all be placed in two orders, the Tabulata
and the Rugosa. The Tabulata appeared first in Chazy times in the
early Middle Ordovician in the shallow seas associated with the Appala-
chian geosyncline of eastern North America, and they remained predomi-
nant throughout the period. The Tabulata may be considered under
six families, Chaetetidae, Calapoeciidae, Syringoporidae, Heliolitidae,
Halysitidae and Favositidae, the first three appearing before the zone
of N emagraptus gracilis in Chazy times, and the last three in Trenton
times, between the zones of Climacograptus peltifer and Climacograptus

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

linearis inclusive. The Rugosa first appeared in two families, Strep-
telasmidae and Stauriidae, in Black River strata (Zone of Nemagraptus
gracilis ), and these two families alone characterise the remainder of
the Ordovician. The number of sub-families under each of these
families is small, at the most three, and the number of genera also is
small.

The Tabulata are always colonial, as are the Stauriidae, but the
Streptelasmidae are predominantly solitary. The Ordovician Rugosa
show no dissepiments and in this respect are more primitive than the
dominantly dissepimented Rugosa of later periods.

The geographical distribution of these Ordovician corals (see text
figs.) is, especially at first, more restricted than in later periods. The
subarctic regions seem specially favoured. The Chazy fauna, which
might perhaps be called the Lichenaria fauna after its dominant genus,
and the Black River fauna, which might similarly be called the
Lambeophyllum-T etradium fauna, seem to be confined to the Appala-
chian seas, with the exception of a possible Black River Tetradium
occurring on Bear Island, south of Spitzbergen. In the Trenton of
N. America and the Upper Caradocian of Britain {i’.e., zones of
Climacograptus peltifer to Pleurograptus linearis inclusive) . corals are
found not only in Appalachian seas but also in the Caledonian seas of
northern Europe and the Himalayan seas of central Asia. The faunas
of all three regions are essentially similar, and we must suppose
relatively free migration from N. America, though it is possible that
the Heliolitidae, Halysitidae and perhaps the Favositidae, which first
appeared during this time, may have originated in Caledonian seas and
spread to N. America. The fauna of all three regions may be called the
Streptelasma fauna from the importance of this genus.

By Cincinnatian times (representing the graptolite zone of
Dicellograptus anceps) a coral fauna had appeared in Tasmania, and
this again is of genera characteristic of Europe and N. America at
that time, indicating free migration. Corals continued richly developed
in N. America and Europe, and the Upper Ordovician or Heliolitid
fauna is a not greatly modified Streptelasma fauna, its most striking
development perhaps being the wealth of Heliolitidae in the
Scandinavian region.

It is to be noted that the present geographical distribution of
Ordovician corals is in the temperate or subarctic regions of to-day and
not at all in the equatorial regions; and this is in marked contrast to
the maximum development in equatorial regions of the reef corals of
to-day.

Facies faunas in Ordovician corals seem to be two (Foerste, 1924,
p. 32). One consists of small solitary non-dissepimented Streptelasma
and perforate stratiform Tabulata like Protar aea ; and this is an obvious
analogue to the Cyathaxonia fauna of the Lower Carboniferous which
in turn is an analogue of the present day coral fauna of cold, deep or
cloudy waters. In Ohio, Indiana and Kentucky, it characterises richly
fossiliferous limestones and clays. The other consists of the compound
Rugosa Favistella, the coenenchymate Tabulata Calapoecia, and the
chaetetid Tetradium. Possibly this is an analogue of the compound or
“reef” coral fauna of the Carboniferous. It characterises the more
arenaceous ( ? clearer water) phases of the Richmond, especially on the
west side of the Cincinnati geanticline, and forms “coral reefs” with
wide lateral but little (a few inches) vertical extent in which the coralla
were not very closely spaced.

THE ORDOVICIAN CORALS.

Morphological Changes. — All Ordovician Rugosa and Tabnlata,
except the later Heliolitidae and Calapoeciinae, are relatively simple
morphologically. The absence of septa in the narrow-celled Lichen-
ariinae, Tetradiinae and Chaetetinae seems a very primitive feature ; the
only variables in this family in the Ordovician are shape of corallum
(in the Tetradiinae, from simple branching to halysitoid, complex
halysitoid and massive hemispherical), and nature of increase, peri-
pheral in Lichenariinae, quadripartite in Tetradiinae and bipartite in
Chaetetinae. The Favositidae have septa, but they are short and equal
and each consists of a vertical series of separate trabeculae (spines) ;
these show no changes during the Ordovician, neither do the tabulae;

‘S>

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

the walls remain unthickened, and there is no interstitial tissue, but the
mural pores are sometimes at the angles and sometimes in the middle
of the walls. The Halysitidae show lability in the shape of the corallites,
which may depart from the usual oval to a polygonal section, and may
form more than one row in the chains; but their tabulae and septa are
like those of Favosites, though only 12 septa are developed. Post-
Ordevician Halysites may develop vertical tubuli between corallites.
The Ordovician Syringoporidae have primitive, sagging tabulae, not
infundibuliform like those of later periods; their septa are like those

THE ORDOVICIAN CORALS.

of Haly sites-, connecting tubules appear by D. anceps time, and varia-
tion in shape, to give prismatic corallites, is present but not in later
periods.

In two families of Tabulata, Calapoeciidae and Heliolitidae,
interstitial tissue develops between tabularia, and greater lability is
shown in these families in the Upper Ordovician than in later times. In
the Calapoeciidae, horizontal canaliculae connecting neighbouring
tabulae develop in horizontal rows between extensions of the septa
which may number 16, 20 or 24 in different species. In the Heliolitidae
the interstitial tissue may consist of small, horizontally based arched
plates; separate trabeculae may develop at right angles to these arched
plates ; or the trabeculae may be so thick as to fill the interstitial space,

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

when they may become arranged in a polygonal spacing. The septa
remain twelve in number and equal, and the tabulae may become highly
domed. In one genus, septa and walls become cribriform.

The Rugose corals of the Ordovician lack dissepiments, the minor
septa remain short, and the tabulae are simple domes. In the Stauriidae
the septa are simple vertical plates, but in the Streptelasmidae the axial
edge is lobed. The Stauriidae show little or no lability, but in
D. anceps time some of the solitary Streptelasmids became triangular
in section; in others the septa developed a wider margin of lobes
axially, and formed an axial structure ; in yet others the septa withdrew
from the axis and lobing became unimportant.

THE ORDOVICIAN CORALS.

The phytogeny of Ordovician corals is a subject of discussion. It
is not difficult, within each family, to work out at least temporarily
satisfactory lines of descent from the oldest species herein assigned to
each family. Thus, in the Chaetetidae, the Tetradiinae and Chaetetinae
could be considered derived from Lichenariinae by a change in the
method of increase, from dominantly lateral or peripheral in Lichenaria
to quadripartite in Tetradium to incomplete bipartite in Chaetetes , the
imperforate and aseptate walls characteristic of the sub-order being
retained throughout. In the Calapoeciidae, the Nyctoporinae may be
presumed with perhaps less certainty to have given rise to the
Calapoeciinae by the development of a characteristic common tissue.
The Syringoporidae may perhaps be considered descended from
Eofletcheria incerta ; though here further evidence from septal struc-
ture is desirable. The Heliolitidae can all be satisfactorily derived
from Protaraea or Coccoseris though Kiaer (1899, 1904) suggests a
different phylogeny, and the Halysitidae from the earliest Halysites
sp. The Favositidae developed naturally from Palaeofavosites. In the
Rugosa, Streptelasmidae can be considered to have developed from
Lambeophyllum by the axial denticulation of the septa. The Stauriidae
seem clearly derived from the earliest Favistella. Weissermel (1897)
considers both Columnaria and Streptelasma to have developed from a
Streptelasma-like ancestor.

When we consider the relation between the various families our
speculations are less satisfactory. Sardeson (1924 fide Okulitch (1935))
considers it possible to derive both Tabulata and Rugosa from
Tetradium; Bassler (in litteris) suggests that Lichenaria is a Rugose
coral and that it may have been the ancestor of all Palaeozoic corals.
One or more of these earliest corals may have arisen from soft-bodied
forms. Possibly the earliest Calapoeciidae, Billingsaria and Nyctopora
arose from Lichenaria by the development of septa, but this does not
seem very likely. There may be a link between the Syringoporidae
and Calapoeciidae; the fasciculate Syringoporid Beuschia was indeed
placed by Kiaer (1930) with the Calapoeciidae. Also, the Halysitidae
may have been derived from Eofletcheria by the assumption of its charac-
teristic mode of increase, and the fixing of the septal number at 12;
possibly the Heliolitidae, with 12 septa, may be derived from
Halysitidae by the development of coenenchyme. But once again these
speculations seem idle without detailed studies. Perhaps the Favositidae
evolved from the Lichenariinae by the development of septa accompanied
by the appearance of mural pores which may have been the result
of crenulation forming at the angles of the walls and becoming pierced
at the apices of the folds ; or they may have derived from the Halysitidae
through Halysites gracilis.

It is difficult to see how the Rugosa Streptelasmidae with two
orders of pinnately inserted septa could have derived from any
Tabulata ; and it is also difficult to see how this family is related to the
Stauriidae, though the common possession of two orders of septa
suggests a relationship.

In the investigation of these early and important phylogenies, our
North American colleagues have the great advantage of the occurrence
within their territories of the earliest Tabulata and Rugosa, and one
looks forward with interest to their future work on them.

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

II. CLASSIFICATION.

Class Anthozoa Ehrenberg 1834 (less Hydrozoa).

Subclass Zoantha de Blainville 1834 (= Order Zoantharia Edwards

1835).

Order Tabulata, Edwards and Haime 1850 (as Sub-order).

Family Chaetetidae.

Family Calapoecjidae

Family Syringoporidae.

Family Halysitidae.

Family Favositidae.

Family Heliolitidae.

Order Rugosa Edwards and Haime 1850 (as Sub-order).

Family Strep telasmidae.

Family Stauriidae.

ORDER TABULATA.

Corallum compound with very slender corallites ; septa short, equal
(except in Nyctoporinae, where two orders are present, alternating),
frequently twelve in number; each a vertical series of spines; walls
often perforate ; tabulae complete or sometimes infundibuliform ;
extratabularial tissue (“coenenchyme”) may be developed.

The Order Tabulata has had many changes in its systematics. As
proposed by Edwards and Haime (1850) it included several genera
which have since proved to belong to other orders, subclasses or even
phyla, such as the scleractinians Pocillopora and Seriatopora, the
hydrocoralline Millepora, the stromatoporoid Labechia , the alcyonarian
Heliopora and the polyzoans Fistulipora, Stenopora, Constellaria ,
Dendropora and Monticulipora. As the true affinities of these misfits
were discovered it came to be thought that there was no such biological
entity as the Tabulata, and the remaining genera were distributed
among the other Anthozoa, chiefly in the Scleractinia and Alcyonaria,
but sometimes in specially named orders (Lindstrom (1876), Zittel
(1879), Nicholson (1879), q.v. for references and historical summary,
Sardeson (1896), Gerth (1908) and Okulitch (1936b)).

The six families listed, with a few more from the later Palaeozoic
rocks, seem to me, however, to form an order clearly distinguishable
from the Alcyonaria, the Rugosa and the Scleractinia, and may well
retain Edwards and Haime ’s name Tabulata. They differ from the
Alcyonaria (except Heliopora) and resemble fhe Rugosa and Selerac-
tinia in having skeletons of fibrous (not spicular as in Alcyonaria)
CaC03, arranged in trabeculae in the vertical skeletal elements and
in sheets in the horizontal plates. They differ from both Rugosa and
Scleractinia in having septa of one order only, and very frequently the
number of septa they contain is 12. They are invariably compound,
the corallites are invariably slender and tend to communicate with one
another by mural pores, connecting tubules, or irregularly perforate
wall tissue and coenenchyma.

The Chaetetidae differ from all others in lacking true septa, any
vertical plates appearing being new walls forming in increase. Also,
the walls are imperforate. Earlier (with Jones, 1940) I regarded the

THE ORDOVICIAN CORALS.

Heliolitidae as a separate order equivalent to the Rugosa ; but later and
wider experience with the Tabulata causes me to lay greater stress on
the similarities of the Heliolitidae to the other Tabulata, rather than on
their differences.

Family Chaetetidae (Tribe Chaetetinae Edwards and Haime
1850).

Massive coralla with very slender aseptate corallites with imper-
forate walls and complete tabulae. M. Ord. — Perm., perhaps also
Jurassic.

The “septa” of many authors (Okulitch, 1935) are not septa as in
the other Zoantharia, but are new dividing walls formed in bipartite,
quadripartite or peripheral increase (Reudemann, 1898).

In Chazy times the Lichenariinae (coralla with unequal increase,
either peripheral or lateral) are dominant; from Black River to Trenton
times the Tetradiinae (with quadripartite increase) gain the ascendency,
and in Richmondian times the Chaetetinae (with incomplete bipartite
increase) appear, the Tetradiinae wane, and the Lichenariinae have
disappeared.

Sub-family Lichenariinae Okulitch 1936.

Increase is unequal but complete, off-sets arising near the margins
of older corallites. M. Ord., N. Amer., Scot.

Lichenaria\ Winchell and Schuchert 1895 (=Lamottia Raymond
1924). Corallites polygonal, erect. M. Ord., N. Amer. Figd.
Winchell and Schuchert, 1895, pi. G, figs. 10-13.

Palaeoalveolites Okulitch 1935. Corallites inclined, upper walls
semicircular. M. Ord., N. Amer. Figd. Bassler, 1932, pi. 7, figs. 2, 3.

Sub-family Tetradiinae Nicholson 1879.

* Increase quadripartite, frequently incomplete. M. and U. Ord.

Tetradium Dana 1848 (—Prismo stylus Okulitch 1935) as for
family. Figd. Bassler, 1932, pi. 21, fig. 19.

Sub-family Chaetetinae.

Increase bipartite, frequently incomplete so that parts of the
corallum are maeandroid. U. Ord.-Perm., perhaps also Jurassic.

Chaetetes Fischer von Waldheim MS. in Eichwald 1829. As for
family. Ord. species figd. Oakley, 1936, pi. 12.

Family Oalapoeciidae nov.

Massive coralla with septa short, thick and spinose, and equal in
number in any species; with interseptal mural pores arranged in
horizontal rows; coenenchyme may be present, formed by extensions
of the tabulae and septa which intersect to enclose tubular spaces
between horizontal floors. M. Ord.-L. Sil.

Chazy and Black River forms are without coenenchyme and may
have 8 longer septa alternating with 8 shorter (Nyctoporinae). In
Trenton times the septa are equal and coenenchyme is developed
(Calapoeciinae).

t For genotypes of, and references to all these genera except ProtrocJiiscolithus
see Lang, Smith and Thomas, 1940 “ Index of Palaeozoic Coral genera.’’ vii +
231 pp., British Museum, London.

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

Sub-family Nyctoporinae nov.

Cerioid coralla with 16 short septa alternating in size. M.-U. Ord.

Nyctopora Nicholson 1879. Corallites with thin walls and septa.

M. -U. Ord., N. Amer., Europe; M. Ord., Asia (Spiti). Nicholson,
1879, pi. ix, figs. 3, 3a-c.

Billing saria Okulitch 1936. Walls thick and septa dilated wedge-
wise. A columella may occur. M. Ord., N. Amer., Europe. Okulitch,
1936, pi. 1, figs. 3, 4.

Sub-family Calapoeciinae.

Cerioid or plocoid coralla; septa equal, 20 or 24. ML Ord.-L. Sil.

Liopora Nicholson and Etheridge 1878. Corallum cerioid. M.
Ord., Europe, N. Amer. Cox, 1936, pi. ii, figs. 3a, 3b.

Calapoecia Billings 1865 (—Columnopora Nicholson 1874;
H ought onia Rominger 1876). Coralla plocoid with coenenchyme more
or less well developed; septa 20, equal. M. Ord.-L. Sil., N. Amer.,
Scandinavia. Cox, 1936, pi. ii, figs. 7-9.

Sarcinula Lamarck 1816 ( = Syringophyllum Edwards and Haime
1850). Corallum alternatingly plocoid and fasciculate; septa equal, 20
or 24. M. Ord.-L, Sil., N. Europe. Cox, 1936, pi. iv, figs. 4-8.

Family Syringoporidae de Fromentel 1861.

Corallum prostrate with corallites conical and anastomosing, or
erect and fasciculate with corallites connected by tubules ; septa
acanthine; wall with fibrous lining; tabulae complete and sagging or
infundibuliform. Ord.-Perm.

In Chazy and Trenton times the corallum is fasciculate, without
connecting tubules ; in the Cincinnatian, connecting tubules appear and
in one genus the corallites become polygonal. The tabulae are complete
and sagging in all Ordovician forms, not infundibuliform as. in
Silurian and later forms.

Sub-family Syringoporinae.

Corallum fasciculate, corallites usually connected by transverse
tubules. Ord.-Perm.

Reuschia Kiaer 1930*. Corallites cylindrical, without connecting
tubules, but with tabulae absent or complete and sagging. M. Ord.,

N. Amer. ; M.-U. Ord., Scandinavia. Kiaer, 1930, pi. iv, figs. 1-3 ;
Okulitch, 1937, pi. xviii, figs. 1-4.

Arcturia Wilson 1931. Corallites polygonal, not in contact, but
with a vertical series of transverse tubuli connecting neighbouring
corallites at each wall angle ; tabulae complete, sagging, not infundibuli-
form. U. Ord., Arctic America. Wilson, 1931, pi. iii, figs. 1-3.

Syringopora Groldfuss 1826. Corallites cylindrical, connected by
transverse tubules; tabulae complete and sagging (in Ordovician
species) or infundibuliform (Sil.-Perm. species). U. Ord., Arctic
America and Rocky Mts. See Troedsson, 1928, pi. 43, figs. 2-3 (for
Ord. species).

* Since this paper went to press, Bassler, 1950, Mem. geol. Soc. Amer. 44: 266
has founded Eofletcheria on Columnaria incerta Billings, 1859. To this new genus
should be removed those American and Scandinavian Middle Ordovician forms with
relatively thin walls and tabulae included in Reuschia above; Reuschia now includes
only thick-walled forms without tabulae, and is known only from the Scandinavian
Upper Ordovician.

THE ORDOVICIAN CORALS.

Family Halysitidae (tribe Halysitinae Edwards and Haime
1850).

Compound coralla with slender compressed corallites united in
chains, each chain typically of one row of corallites, the ends of the
chain being connected to the sides of others; smaller tubules crossed
by tabulae may occur between the corallites; walls imperforate, septa
acanthine, 12 in number, equal; tabulae complete, sagging. M. Ord.-
Gedinnian. Ordovician species show no tubuli between corallites.

Holy sites Fischer von Waldheim 1813 {—Catenipora Lamarck
1816). Corallites round or oval in transverse section, occasionally with
more than one row in a chain when they become polygonal; smaller
tubules between corallites rare or absent in Ordovician species.

M. Ord.-Sil. ; Gedinnian in Asia Minor. Lambe, 1899, pi. iii, figs. 5-7.

Labyrinthites Lambe 1906. Corallites prismatic, thick walled, in
extremely short chains, frequently with two rows of corallites in a
chain; no interstitial tubules. Ord., Arctic America. Lambe, 1906,
p. 328, text fig.

Family Favositidae Edwards and Haime 1850.

Massive coralla without coenenchyme ; with slender prismatic
corallites with mural pores; septa short, equal, spinose; tabulae
complete. U. Ord.-Perm.

Early upper Ordovician forms have the mural pores at the angles
of the walls ( Palaeofavosites ) ; towards the top of the Ordovician these
are joined by forms with pores in the middles of the walls.

Sub-family Favositinae.

Corallum with prismatic, thin-walled corallites. U. Ord -U. Dev.

Palaeofavosites Twenhofel 1914. Mural pores occur predominantly
at the angles of the walls. U. Ord.-M. Sil. Lambe, 1899, pi. i, fig. 2.

Favosites Lamarck 1816. Mural pores occur predominantly in the
middle of the walls. U. Ord.-M. Dev. Lambe, 1899, pi. i, fig. 1.

Family Heliolitidae Lindstrom 1876.

Massive coralla with slender corallites separated by coenenchyme,
each corallite with 12 equal spinose septa and complete tabulae. M.
Ord.-U. Dev.

The earliest Heliolitidae (Trenton) have thick trabeculae and little
coenenchyme and some may have perforate septa; upper Ordovician
forms are very numerous and diverse.

Sub-family Coccoserinae Lindstrom 1899.

Encrusting or discoid coralla with coenenchyme almost absent or
tubular, the tubuli being closed or restricted by thickening of the
vertical trabeculae forming their walls; each corallite filled or almost
filled by thick septal trabeculae which curve upwards and inwards to
become vertical at the axis. M. Ord.-L. Sil., Europe, N. Amer., Aust.

Protaraea Edwards and Haime 1851 ( =Stylaraea Seebach 1866
non Edwards and Haime 1851, Tumularia Robinson 1916). Coenen-
chyme very narrow or absent, trabeculae very stout. M. Ord.-L. Sil.,

N. Amer., Europe, and Aust. Foerste, 1924.

Coccoseris Eichwald 1855. Coenenchyme wide; trabeculae so stout
that all spaces are filled. U. Ord.-L. Sil., Europe. Lindstrom, 1899,
pi. xii, figs. 3, 4, 6, 7.

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

Acidolites Lang, Smith and Thomas 1940 (=Acantholithu$
Lindstrom 1899 non Stimpson, 1858). Coenenchyme wide, tubular;
trabeculae relatively slender, leaving spaces crossed by tabulae. M.
Ord.-L. Sil., Europe, Tasmania. Lindstrom 1899, pi. xi, figs. 23, 24,
26, 30.

TrochiscoUthus Kiaer 1899. Corallum branching; coenenchyme
tubular, moderately wide; axis of branch with slender trabeculae
forming occasionally perforate septa and walls; outer part of branch
with all spaces filled by thickening of the trabeculae. M. and U. Ord.,
Europe. Kiaer, 1904, fig. 5-6, 3.

Protrochiscolithus Troedsson 1928. Coenenchyme very narrow;
trabeculae slender, forming occasionally perforate walls and leaving
spaces in the corallum. U. Ord., Arctic N. Amer. Troedsson, 1928,
pi. 30.

Urceopora Eichwald 1855 and Stylidium Eichwald 1855 from the
Baltic are possibly Coccoserids, but thin sections are required.

Sub-family Proporinae nov.

Massive cor alia with slender tubular corallites separated by a
coenenchyme of domed plates and isolated trabeculae ; septa 12, spinose,
equal. M. Ord.-U. Sil.

Propora Edwards and Haime 1849 ( Lyellia Edwards and Haime
1851, Pinacopora Nicholson and Etheridge 1878, Nicholsonia Kiaer
1899 non Schluter 1885). Thickening of trabeculae not greater in
peripheral or surface zones than in axial or lower zones of the corallum ;
tabulae flat or sagging. M. Ord.-U. Sil. Lindstrom, 1899, pi. viii,
figs. 8-10.

Plasmoporella Kiaer 1899. Trabeculae very slender; tabulae domed
and complete or incomplete. U. Ord.-M. Sil., Europe, Tasmania, and
N. Amer. Kiaer, 1899, pi. v, figs. 9, 10.

Diploepora Q;uenstedt 1879. Corallum branching; axial part of
branch with slender trabeculae ; peripheral part with all spaces filled
by thickening of the trabeculae. U. Ord.-Sil., Europe. Kiaer, 1899,
pi. v, fig. 7.

Sub-family Proheliolitinae Kiaer 1899.

Massive coralla with slender corallites in contact or polygonal or
separated across their angles by one, two or three small coenenchymal
tubules, each of which may increase in diameter to form a normal
corallite; septa 12, equal, each represented by a single vertical series
of very short, downwardly directed, discrete spines. U. Ord.-L. Sil.,
Europe.

Proheliolites Kiaer 1897 as for sub-family. Kiaer, 1899, pi. iii,
figs. 5, 6.

ORDER RXJGOSA.

Solitary or compound coralla in which, after the insertion of the
first six septa, meta-septa are inserted at four points only. Typically
there are two series of septa and a varying development of tabulae
and dissepiments. The corallum invariably has an epitheca. Compound
coralla are formed by axial, peripheral or lateral increase. M. Ord.-
U. Perm. Ordovician Rugosa have no dissepiments.

THE ORDOVICIAN CORALS.

Family Streptelasmidae Nicholson 1889.

Solitary conical Rugose corals with short minor septa and without
dissepiments ; the major . septa are denticulate axially ; early septal
dilatation decreases giving a peripheral stereozone only in adults;
tabulae domed, complete or incomplete. M. Ord.-M. Sil.

The width of the axial zone of denticulation of the septa generally
increases in Ordovician forms; in the Cincinnatian many species
become angulate (especially triangular) in transverse section.

Lambeoplvyllum Okulitch 1938. Septal denticulations few or
absent ; no axial structure ; in adult stages the axial edges of the major
septa of the cardinal quadrant run together to make a wall for each
side of the cardinal fossula which contains the long cardinal septum,
and the axial edges of the septa in the counter quadrants run together
to form a wall along the counter sides of the alar fossulae. M. Ord.,
N. Amer., Baltic. Okulitch, 1938, p. 99, text fig. B.

Streptelasma Hall 1847. Septa long, axial denticulations few and
axial structure narrow. M. Ord.-Sil. Cosmop. Cox, 1937, pi. i, figs. 1, 2.

Grewingkia Dybowski 1873 ( =Kiaerophyllum Wedekind 1927).
Septa short, axial denticulations numerous, axial structure wide and
present in young stages. U. Ord., N. Amer., Europe. Wedekind, 1927,
pi. i, figs. 8, 9.

Bracinyelasma Lang, Smith and Thomas 1940 ( =Dybowskia

Wedekind 1927 non Dali 1876). Axial denticulations few in young
stages ; in adult stages without axial structure and with septa withdrawn
from the axis. U. Ord., Europe. Wedekind 1927, pi. i, figs. 10, 11.
Scheffen, 1933, pi. i, figs. 5, 7.

Palaeophyllum Billings, 1858. Phaceloid Streptelasma. M.-U. Ord.,
Canada. Lambe, 1901, pi. vi, fig. 3, 3a, b.

Holophragma Lindstrom 1896. Calceoloid with cardinal septum
on flattened side longer than other septa, but not produced upward
into a columella; septa long, so thick as to fill all interseptal loculi
and without axial denticulations. ? U. Ord., N. Amer. ; Sil., Scandinavia.
Lindstrom 1896, pi. vi, figs. 77, 78, 81. Possibly a halliid.

Coelostylis Lindstrom 1880. Like Streptelasma but with a blade-
like columella produced upwards from the axis. M. Ord., Scandinavia.
Lindstrom in Angelin, 1880, pi. i, fig. 11.

Family Stauriidae Edwards and Haime 1850.

Compound Rugose corals with small corallites; the minor septa
may be short and dissepiments absent, or longer when lonsdaleoid
dissepiments may develop. Major septa long, axial edges smooth, no
axial structure; tabulae complete, flat or funnel-shaped. M. Ord.-M,
Dev. The Ordovician forms are all without dissepiments.

Favistella Dana 1846. Fasciculate or cerioid Rugosa with small
corallites with short septa and without dissepiments; tabulae complete
and nearly horizontal; walls thin. M. Ord.-L. Sil., cosmop. Lambe,
1901, pi. vi, figs. 1, la.

III. SEQUENCE OF FAUNAS.

North America.

Chazy ( pve-Nemagraptus gracilis zone).

The lower Middle Ordovician Chazy corals may well be the oldest
in the world. A record of an earlier occurrence (Tetradium? simplex
Bassler (1919)) refers to such poorly preserved material that it is

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

doubtful to what phylum it belongs. The Chazy fauna seems to be
older than the N emagraptus gracilis zone in the graptolite sequence
(though possibly of it) since recent wTork (Cooper and Cooper, 1946)
shows that in Virginia the beds containing it (Newmarket limestone)
lie below the Edinburg formation with Normanskill graptolites such as
N. gracilis and Black River and perhaps early Trenton shelly faunas.

The Chazy corals are all compound and seem all to be Tabulata.
Two genera are placed here in the Chaetetidae, one in the Calapoeciidae
and the fourth somewhat doubtfully in the Syringoporidae. They are
known as follows, from north to south. [In this paragraph, as in all
others in this paper citing the occurrence of genera and species in
formations, I have relied on figures of thin sections, or on thin sections
in the Sedgwick Museum, the British Museum (Natural History) or
the University of Queensland; records of Ordovician coral species are
in need of precise revision, the names usually being a guide only to the
external form or unsectioned appearance of the fossil. Formations are
cited so that possible errors in correlation may be rectified.] Mingan
formation, Mingan I., Billingsaria parva (Bill.) and Eofletcheria incerta
(Bill. Lambe 1899 and Twenhofel 1938) ; Aylmer formation,
Ottawa valley, the single-tubed Tetradium cylindricum Wilson (1921)
which according to Okulitch (1935) is conspecific with the Black River
T. spring op oroides Ulrich; Chazy of L. Champlain region, B. parva ,
Lambe (1899) ; upper part of lower Chazy (Day Point division) of
Vermont, Lichenaria ( Lamottia ) lieroensis Raymond (1925) and at
Plattsburg, N.Y. ; Newcastle limestone of Virginia, B. parva, Cooper
and Cooper (1946) (records only) ; Lenoir of E. Tennessee, Schuchert
(1943), B. parva, Lichenaria prima Okulitch (1936). The Stones River
group of Tennessee and its correlatives in neighbouring states, from
which Okulitch (1936) listed Rugose corals, are now regarded as Black
River rather than Chazy ( Schuchert ( 1943 ) ) .

Lamottia and Lichenaria seem to me to be congeneric, though
Okulitch (1936) states that perforations occur in the walls of Lamottia
and not in Lichenaria ; topotype specimens of Lamottia I have examined
at the Sedgwick and British Museums do not show perforations.
Lichenaria forms massive coralla of polygonal tabulate tubules, aseptate
and aporose as in the Chaetetidae, in which I place it ; increase is
apparently peripheral and indicates a sub-family separate from the
Chaetetinae and Tetradiinae which I also regard as Chaetetidae.
Tetradium is aseptate and aporose with tubules 4-sided, but in this early
age its species is single-tubed ; each tube divides characteristically by
quadripartite fission, the new walls growing inwards from the middles
of the old walls. This inw^ard growth of the new walls is seen in the
bipartite or incomplete fission of typical chaetetids. Okulitch (1939)
has suggested how the Black River, Trenton and Richmond tetradiids
may have developed from this single-tubed form by an increase in the
rate of fission producing at first chain-like or fasciculate coralla, and
later cerioid massive coralla.

Billingsana differs from all other Palaeozoic corals except the later
Ordovician Nyctopora in possessing the fixed number of 8 long
( ? lamellar) septa alternating with 8 short septa. It differs from
Nyctopora by the wedge-like thickening of its septa and by its columella;
its corallites communicate by irregular vacuities between the trabe-
culae. These two genera I regard as forming a distinct Ordovician sub-
family, the Nyctoporinae, and there are grounds for believing that it

THE ORDOVICIAN CORALS.

and the Calapoeciinae of the later Ordovician are related. A thin section
of Nyctopora billingsi Nich. in the British Museum from the Trenton
of Nepean Pt., Ottawa, shows a horizontal row of wall pores as in
Calapoecia.

Fasciculate coralla, whose tubules are slender (about 1 mm.) and
aporose, and whose possession of spinose septa is doubtful (compare
Lambe (1899) and Okulitch (1937)), but which are crossed by tabulae
(occassionally infundibuliform) are assigned to the family Syringo-
poridae. Bassler (1950) has founded the genus Eofletcheria for them.

Nemagraptus gracilis zone.

(Black River.)

As with the Chazy, the relation of the Black River group of the
type area to the standard graptolitic succession rests on indirect
evidence. The correlation of N. American strata with those of the Black
River of the type section in New York State has recently been revised,
so that the assessment of Chazy and Black River faunas made by
Okulitch (1936, 1938) requires modification. For instance the Stones
River formation of Tennessee and its correlatives in neighbouring
states are now regarded as Black River, not Chazy ( Schuchert ( 1943 ) ) ,
and the Cloche I. limestone and correlative beds at Pauquette’s Rapids
in the upper Ottawa Valley are regarded as early Trenton, not Black
River (Kay (1937)). The Edinburg formation of Virginia has been
found by Cooper and Cooper (1946) to contain shales with graptolites
of the Nemagraptus gracilis zone intergrading with and laterally
equivalent to limestones, etc., with shelly faunas of Black River type,
so that it may be deduced that the type Black River is also of the
N emagraptus gracilis zone.

In the area of continuous Black River outcrops from New York
into S.E. Ontario (Young (1943)) which contains the type area,
Billingsaria and E. incerta seem to have disappeared, though Lichenaria
is still present. Tetradium is now common, being represented by several
species in which the rate of fission outsteps the rate of branching, so that
thick branched ( T . cellulosum), halysitoid ( T . halysitoides) or even
hemispherical cerioid coralla (T. fibratum) develop alongside T. syringo-
poroides Okulitch (1938, 1939). But the outstanding feature of the
Black River fauna is the presence for the first time of Rugosa, the Order
in which septa (major and minor) are inserted in only 4 positions in
the corallum. The earliest genera seem to be solitary corals — strep-
telasmids — Lambeophyllum Okulitch (1938) and f Coelostylis

Lindstrom (1880), (i.e., “ Lindstroemia” whiteavesi Foerste (1906),
description only, no figure available) ; the latter is columellate, one
septum being elongate and swollen axially. In addition, cerioid
coralla occur which are usually referred to (i Columnaria” ; but
specimens from the type area of Black River strata seem not to have
had their internal structure sufficiently figured or described for us to
be certain whether they are Rugosa or not. The records usually read
Columnaria halli, or Columnaria alveolata, but the specimens need
careful study; many may be Favistella ; some may be Lichenaria, e.g.,
Columnaria simplissima Okulitch. The specimens figured by Lambe
(1901) from fault blocks in Renfrew Co., Ontario, seem to be Rugosa,
but may well belong to differing genera, and also quite possibly came
from early Trenton rather than Black River strata. Calapoecia and
Streptelasma have both been recorded from a fault block at Pauquette’s

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

Rapids, upper Ottawa valley, in strata previously considered Black
River but now (Kay (1937) ) referred to early Trenton, so that it seems
that these two genera may not now be included in the Black River
fauna.

In the States south and south-west of New York, e.g., in Tennessee
in the Stones River group and its correlatives in neighbouring states,
and in the Decorah Formation of Minnesota, there are additional
endemic forms. These are the insufficiently described small Rugose cup
coral ? Streptelasma parasiticum Ulrich (Decorah of Minnesota and
Lebanon of Tennessee (Okulitch (1938)), and the compound, cerioid
Rugosa “Columnaria” minor Bassler from the Kimmswick formation
of Tennessee (specimen in B.M.), the colonial Tabulata Palaeoalveolites
carterensis with reclined corallites and Lichenaria carterensis both
from the Carters limestone of central Tennessee (Okulitch (1938)), and
L. typa from the Decorah of Minnesota. Okulitch suggests that
Palaeoalveolites has mural pores and septal spines and that it is
ancestral to the Silurian Alveolites, but its structure and affinities
require elucidation. It could well be a development from Lichenaria.
Nyctopora apparently makes its first appearance (specimen in B.M.)
in the “ Murat” of Virginia; Cooper and Cooper (1946) regard this as
post-Chazy, and either pre-Black River or early Black River.

Post-A. gracilis to Pleurograptus linearis zone.

(Trenton.)

The Trenton group is of considerable extent in N. America, the
standard now used (Kay (1937) ) extending in Ontario from the base of
the Rockland formation to the top of the Gloucester shale ; it is roughly
equivalent to the three graptolite shales Magog, Canagoharie and Utica,
covering the American graptolite zones from Cyrtograptus tricornis
insect if or mis to Glossograptus quadrimucronatus typus inclusive which
in the British sequence probably represent the zones above Nemagraptus
gracilis to and including Pleurograptus linearis (Thorslund (1940)).

The Trenton coral fauna comes almost entirely from the pre-Utica
(i.e., pr Q-linearis zone) section of the Trenton group. At Pauquette’s
Rapids in a fault block in the upper Ottawa valley a fauna occurs
which was previously considered Black River but is now (Kay (1937)),
regarded as Rockland. It is distinguished by the first occurrence of
Calapoecia (as C. canadensis ) and of Streptelasma (as S. corniculum)
together with the insufficiently known Rugose “Columnaria” magnifica
Okulitch (1938), a phaceloid species with large corallites, the solitary
Lambeophyllum ? apertum (Bill.) Okulitch (1938), and the curious
Palaeoalveolites pauquettensis Okulitch (1938), while Lambe (1901)
records also “Columnaria” halli Nich., Lambeophyllum profundum
and Tetradium fibratum, and Okulitch (1938) records Lichenaria typa.
Calapoecia shares with Liopora and Syringophyllum the possession of a
single order of septa, constant in number in the corallites of any one
species, and of communicating canals between corallites developed
between the septa periodically at the same heights in neighbouring
coralla.

From the slightly younger typical development in Nbw York
(Trenton) and Ontario (Peterborough), Nicholson (1875) described
a small fauna, “Columnaria alveolata,” Streptelasma corniculum and
massive Tetradium as in the Rockland formation, with in addition
Nyctopora , and the earliest Heliotidiae from N. America. These are
Protaraea vetusta Hall and Propora goldfussi (Bill.) with 12 equal septa,

ORDOVICIAN CORAL£>.

though Trenton specimens of the latter typically Richmondian species
have not been figured. Foerste (1924) remarks that the Trenton
Protaraea were unattached except for a small central point, while the
later, Richmond forms were encrusting. Both these species are known
also from Scandinavia. The Scottish calapoeciid Liopora is represented
(specimen in B.M.). The Trenton fauna is widespread in N. America.
In the Hermitage shale of Tennessee which Schuchert (1943) considers
to represent the lower part of the Trenton group, typical species are
accompanied by Lichenaria, Nyctopora crenulata Bassler (1932)
showing lamellar septa, and by Aulopora, the latter recorded by Bassler
but not figured. If this Aulopora is correctly identified it forms the
earliest representative of the genus, and the upper Ordovician
Syringoporidae may well have developed from it, rather from Beuschia
as previously suggested.

An interesting first occurrence for America is Halysites, from the
upper part of the Liskeard formation of L. Timiskaming, Out., which
Kay (1937) correlates with the Coburg limestone and the early part of
the Utica shale (P. linearis zone). Teichert (1937) has recorded H.
aequabilis from possibly upper Trenton strata on Iglulik I. in Arctic
N. America, together with Propora lambei and Coelostylis f oppletus,
a solitary Rugose coral with septa highly dilated throughout, their axial
parts forming a raised columella in the floor of the calice.

Dicellograptus anceps zone.

Cincinnatian.

The Cincinnatian of N. America (Schuchert (1943)) saw a con-
siderable development of strata; in the typical Ohio-Indiana-Kentucky
sequence the Eden and Maysville formations at the base are followed
by the Richmond ; the two first are usually regarded as equivalent to the
Lorraine of New York and the Hudson river group of Vanuscem (1842)
and Hall (1847). If the Utica shale is correctly correlated with the
British Zone of P. linearis (Thorslund (1940)), then the Cincinnatian
may be roughly equivalent to the Ashgillian, i.e., to the Dicellograptus
anceps zone of the British Isles.

Corals do not occur in the Eden or Maysville formations, or in the
Lorraine, but are common in the Richmond. In Ohio, Indiana and
Kentucky, Foerste (1909) gives a Richmondian assemblage of Strep-
telasma, some solitary and like the Grewingkia of Europe, some like
“ S. divaricans Nich.”, conical individuals aggregated by their bases,
some “ Columnaria” (various cerioid and phaceloid forms, insufficiently
illustrated) with Calapoecia, Tetradium of cerioid habit, and encrusting
species of Protaraea.

In Richmond equivalents in Ontario and Quebec, Foerste (1924)
figures a similar fauna with Propora goldfussi added. Palaeophyllum
rugosum Bill., which, according to Lang, Smith and Thomas (1940) is
a phaceloid Strepelasmid, is possibly from Richmond strata on L. St.
John (Cox (1936)), though Dresser’s (1916) map indicates a Trenton
age.

It may be that the Red River formation of Manitoba is older than
Richmondian; its fauna generally resembles that cited above, but in
addition it contains Halysites gracilis in which each chain of corallites
may consist of more than one row of corallites (Leith (1944)), while in
its upper layers Palaeofavo sites makes its first appearance in America

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

as P. prolificus (Foerste (1929) recorded only). In the Stony Mountain
formation above the Red River, which seems a Richmond correlative
(Okulitch (1943)), the large streptelasmids are angulate ($. trilo-
batum), some of the Streptelasmidae are columellate and calceoloid
( Holophragma anticonvexa Okulitch), Halysites and Calapoecia are
associated with Palaeofavosites and, according to Okulitch, with Favosites
intermedins , with characters said to be intermediate between those of
Palaeofavosites and Favosites. This would seem to be the first occurrence
in America of the Favositidae.

In the N. American Arctic regions (including Alaska) several
Richmondian (or possibly Red River) correlatives have been recorded
(Cox (1937), Teichert (1937), Roy (1941), Wilson . (1931) ). Strep-
telasma rusticum occurs with angulate (particularly trilobate) forms and
columellate ? Coelostylis and Holophragma. “Columnaria” is recorded
(Troedsson (1928)), Calapoecia is common, particularly C. anticos-
tiensis, with Halysites , Propora (often listed as Plasmopora lambei ),
Syringopora with tabulae which are not infundibuliform (Miller and
Youngquist (1947)), and the curious syringoporid with polygonal
corallites, Arcturia Wilson (= Labyrinthites Troedsson non Lambe).
Labyrinthites Lambe is a curious form, possibly related to Halysites,
recorded from C. Chudleigh in Hudson Str. In the probably Rich-
mondian (Red River) C. Calhoun beds of W. Greenland, the heliolitid
Protrochiscolithus occurs (Troedsson (1928)). Chaetetes has been
described (Oakley (1936)) and this is possibly the earliest authentic
occurrence for a typical member of this long-ranging genus. Tetradium
seems to be absent.

The Richmond correlative in the Rocky Mountains, the Beaverfoot
formation (Wilson (1926)), has large solitary Streptelasmidae resem-
bling Grewingkia, some being angulate and trilobate (8. prolong at um) ;
others have a columella ( ? Coelostylis patellum) . Cerioid and phace-
loid “Columnaria” (insufficiently figured) occur, and an interesting
feature is the phaceloid Rugosa assigned by Wilson to Diphyphyllum.
These are insufficiently known but appear to represent a new endemic
Rugose element in the coral fauna. Syringopora has entered here, as
in the Arctic regions, with concave, not infundibuliform tabulae,
associated with Calapoecia, Halysites without interstitial tubules,
Palaeofavosites and species unfigured but referred to Favosites. Tetra-
dium seems to be absent.

On Anticosti Island (Twenhofel (1927)), the English Head and
Vaureal formations have a Richmondian fauna, with Streptelasma
rusticum and large angulate Streptelasma, (cf. Cox (1937)), “Colum-
naria” cerioid and fasciculate and insufficiently illustrated, Calapoecia ,
Halysites, Propora goldfussi (Bill.) and Palaeofavosites. The Ellis Bay
formation is considered by Twenhofel to be post-Richmondian
(Gamachian) and indeed its coral fauna contains at least one new
element in “ Cyathophyllum” ellisense Twenhofel (1927), in addition to
the characteristic Richmond genera. This new form suggests the Silurian
Strombodes and Pilophyllum. Aulopora and Favosites also occur in the
lists, but no sections are figured. Tetradium seems to be absent from
Anticosti.

British Isles.

Craighead Limestone ( ? Climacograptus peltifer zone).

The Craighead limestone of Girvan, Scotland, would now seem in
all probability to belong to the Climacograptus peltifer Zone. Anderson

THE ORDOVICIAN CORALS.

and Pringle (1946) have traced the field relations between fossiliferous
olive-green sandy mudstones and limestones as the Craighead Quarry
has been extended, and have concluded that the Craighead limestone is
a reef phase in these mudstones. As they consider the mudstone fauna
to be that of the Balclatchie mudstones, they regard the contemporaneity
of the Craighead limestone and Balclatchie mudstones as established, so
that the Craighead limestone is younger than the Stinchar limestones.
They conclude that the horizon of the Craighead limestone is thus at the
top of the Glenkiln shales or the base of the Hartfell, with a slight
balance in favour of the former. Bulman {verb, diet.), who has recently
(1944-47) studied the Balclatchie graptolites of Laggan Burn, considers
these to be not younger than the zone of Climacograptus peltifer , and
not older than the zone of N emagraptus gracilis, and that they are pro-
bably of the former zone, i.e., early Caradocian, upper Glenkiln. The
Craighead limestone, which contains the oldest coral fauna in the British
Isles, is therefore not so old as the Chazy of N. America, but could be
coeval with some part of either the Black River or the early Trenton,
probably early Trenton ; and it is interesting to note therefore that its
assemblage of genera is of Trenton rather than Black River or Chazy
aspect.

Its Rugosa are probably all Streptelasmidae. Streptelasma craig ense
Nicholson and Etheridge (1878) is the commonest. S. fossulatum Wang
(1948) could well be Lambeophyllum , the typical N. American Black
River genus, which seems to persist into early Trenton (Rockland)
times. “Lindstroemia subduplicata McCoy” N. and E. is not columellate
like “Lindstroemia” ( ? Coelostylis ) whiteavesi Foerste from the Ottawa
Black River, but suggests relationship to Streptelasma aequiseptatum
McCoy of the Robeston Wathen and Coniston limestones. S. aggregatum
is fasciculate. No “ Columnar ia,” so characteristic of the N. American
Middle and Upper Ordovician, occurs, and in this particular this
Scottish fauna presents a strong contrast to the North American.

The Nyctoporinae are represented by Billingsaria occidentals,
though in N. America this genus apparently died out at the end of
Chazy times. The Calapoeciinae are represented by Liopora favosa
which in its structure is quite similar to the Trenton L. americana
Bassler and to Calapoecia canadensis.

As in the Trenton of N. America, the Heliolitidae and Halysitidae
are present. The only heliolitid is an irregularly ramose form of very
thickened tissue, wrongly referred by Nicholson and Etheridge (1878)
to Heliolites grayi Edw. and H., but which Kiaer (1932) suggests may
be Trochiscolithus. This form may however be Protaraea. The
Haly sites seems to be the oldest so far recorded; several specimens are
in the British Museum.

The Chaetetidae are represented by only one species of Tetradium,
of a massive type such as characterises the Trenton in N. America ;
true Chaetetes seems absent, for Chaetetes sp. Nich. and Eth. (1878)
is probably polyzoan.

Whether the Favositidae are already present in the Craighead
limestone is very doubtful. Nicholson and Etheridge (1878) described
Favosites girvanensis, but this is apparently without mural pores or
septa, though its walls are crenulate at the angles; it may be referable
to the N. American Middle Ordovician Lichenaria. The peculiar
“ Thecostegites f” scoticus N. and E, is difficult to place. Its eorallites

20 PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

are polygonal, with thickened walls, apparently aporose, and open to the
surface obliquely suggesting an irregularly dendroid development with
thickened walls from the aporose Lichariinae, after the manner of
Thamnopora from the later Favositidae.

In its general character, therefore, the Craighead fauna is similar
to the early Trenton. Thus Streptelasma is present in association with
Lambeophyllum. The Calapoeciidae are represented by Liopora. The
Heliolitidae are represented by forms of dense structure only; the
Halysitidae are present. The Lichenariinae are present, but crenulation
of the walls is very marked in the only species. Tetradium is present
as a massive species only. The differences between the two are also
striking; thus Favistella is absent; and Billingsaria, not Nyctopora, is
the characteristic nyctoporid.

Robeston Wathen Limestone.

(? Zone of C. wilsoni or Dicranograptus clingani.)

The next youngest coral fauna in the British Isles seems to be
that collected from the Robeston Wathen limestone of S. Wales
(Haverfordwest), which, according to Pringle and George (1948, p.30),
is developed at about the horizon of the vulgatus beds some 90 feet
below the top of the Mydrim shales. The assemblage of graptolites in
the vulgatus beds perhaps suggests the C. wilsoni or the D. clingani
zone of Scotland, but Jones (1936) states that the limestone occurs
in the ^horizon of Mesograptus multidens, so that it may be older,
perhaps in C. peltifer zone. It appears younger than Nemagraptus
gracilis zone, for the Mydrim limestone at the base of the Mydrim shales
has a Nemagraptus gracilis assemblage.

The Fron Qy. fauna in the Sedgwick Museum consists of
Streptelasma cf. aequisulcatum, Holy sites (2 species) and the heliolitids
Propora hirsuta Nichols onia” megastoma (McCoy), which also
occurs in Fx at Soida, Estland) and Propora sp. nov., with very large
tubules. Also from the Robeston Wathen limestone of unspecified
locality are the streptelasmid Brachyelasma (which occurs in 5a of
Norway), a doubtful Syringaxon (which, if it could be substantiated,
would be the oldest representative of a long-ranging genus), and
Trochiscolithus ? inordinatus (Lonsdale) a slenderly branching
heliolitid with very dense tissues. Syringophyllum is represented by a
single specimen from Grondre; this calapoeciid genus is not known in
N. America, just as Calapoecia itself is not known in the British Isles,
though the two occur together in Norway (5a).

This fauna, with the occurrence of Propora and Syringophyllum ,
has a younger aspect than the Craighead. The absence of Tetradiinae,
Stauriidae, Lichenariinae and Palaeofavo sites is notable.

Coniston Limestone.

(? Zone of Pleurograptus linearis or Dicellograptus anceps.)

This term “ Coniston Limestone” originally included the few
hundred feet of marine sediment between the Borrowdale Yolcanics
and the Silurian Stockdale shales in the north of England, but has
lately (Eastwood (1935) ) been restricted to the lower 100 feet, which
Marr (1916) regarded as Caradocian. The corals from the upper 100
feet are mostly from the White Kiesley and equivalent limestones,
usually correlated with the Dicellograptus anceps zone. King and
Williams (1948) have suggested that the Coniston Limestone as at

THE ORDOVICIAN CORALS.

present restricted may represent part of the D. anceps zone also,
but the evidence is indirect, and most authors have regarded it as older
than the D. anceps zone and possibly of the zone of P. linearis.

From the Museum collections available it does not seem possible
to be sure which species of coral are confined to Marr’s Caradocian
and which to his Ashgillian, with the exception of those from the
Ashgillian Kiesley limestone; but, in the main, specimens seem to be
from the Sleddale group. Spring ophyllum organum is abundant, as is
Streptelasma aequisulcatum and Holy sites. These three have some
resemblance to the Robeston Wathen fauna, as does a wide-tubed species
of Propora. Other species of Heliolitidae, e.g. Propora cf. goldfussi
(Bill.) (S.M.F. 15224, Helm Gill, Dent) and calical moulds referred
to (( Heliolites tubulatus and subtubulatus” are different from those of
Robeston Wathen. The most striking difference from Robeston Wathen
however is the occurrence of Palaeofavosites sp. at Crag Hill, Yorks,
and of P. crassa at Coniston Waterhead.

Portraine Coral Bed.

The Irish Portraine Coral Bed which possibly is equivalent to
the Coniston Limestone (restricted to Sleddale Stage) has an important
fauna which, however, is mostly unsectioned. Propora cf. goldfussi
(Bill.) occurs as in the Coniston fauna.

Ashgillian (Zone of D. anceps, which includes developments
of the dwarf D. complanatus fauna (Elies (1937) ).

The Ashgillian shelly faunas of North Wales (Denbighshire) have
a fair coral fauna which unfortunately has not been investigated by
thin section. Spring ophyllum, Palaeofavosites, Halysites, Propora

(with wide tubules) and external moulds of Heliolitidae, some of which
suggest Proheliolites dubius or Propora cf. goldfussi (Bill.), are
represented in the B.M. collection from Mynydd Fron Frys and nearby.

The Ashgillian Kiesley limestone of the Lakes District contains
Palaeofavosites, Halysites and Kiaerophyllum.

Bear Island.

(? Zone of Nemagraptus gracilis.)

On Bear I., off the north of Norway, Tetradium spring oporoides
has been found in strata correlated by Holtedahl (1918) with the Black
River of N. America, which is reasonably regarded as of Nemagraptus
gracilis zone. Holtedahl remarks that the associated fauna is purely
American, showing no relation to the Middle Ordovician of N.W. Europe.

Norway.

Kalstad Limestone ( ? Zone of D. cling ani).

In Norway, one of the oldest coral faunas occurs in the Kalstad
limestone in Meldalen. Its stratigraphical position is not satisfactorily
known. Kiaer (1932) considered its coral fauna compared well with
that of the Mj^s limestone in the Oslo district, which is now usually
(St^rmer, 1945) regarded as between zones 4b 3 and 4c a, i.e., the
D. cling ani zone. Vogt (1945) says that it could be equated with the
Svattgern rather than with the Holonda of Meldalen; if it correlated
with either, the outside limits for its age should be (from stratigraphical
relations) the Lower Llanvirn and Middle to Upper Caradoc ( Dicrano -
graptus shale, i.e. zone of D. clingani). Kiaer also compared it with
the Craighead limestone which as seen above is possibly C. peltifer zone.

22 PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

The fauna seems very like that of the Trenton of North America.
Thus, of the Rugosa, Streptelasma occurs, and a cerioid form which
Kiaer has placed in Columnaria, remarking however that it shows no
close resemblance to the N. American ‘Columnaria’ and may indeed be
a cerioid Streptelasma. There is also a curious solitary coral, quite
unlike any other Ordovician Rugosa in that it has a rich wide
dissepimentarium and very numerous carinate septa, the minor septa
being long and leaving only a narrow tabularium in which the major
septa extend to the axis (0. kjerulfi).

The Halysitidae and Heliolitidae are both represented, the latter
by a Propora cf. goldfussi (Bill.) with resemblances to a form from
the Robeston Wathen and Coniston limestones of England. A small
cerioid form has been placed doubtfully in Nyctopora by Kiaer ;
however, it could well be Liopora. Both these genera are common in
the Trenton. A small fasciculate form, much recrystallised, is regarded
by Kiaer as Liopora incerta (Bill.), but, until its minute structure is
better known, we cannot be certain whether it be Syringoporidae,
Calapoeciidae or Rhabdocyclidae. No Lichenaria or Tetradium , so
common in the Trenton of N. America, is present; nor Palaeofavo sites
such as characterised the Richmondian and Ashgillian D. anceps zone
and perhaps also the P. linearis zone in N. America and the British
Isles.

The Mj</>s limestone, which seems to be similar, has the fasciculate
“Liopora incerta,” the cerioid Liopora favosa and PLaly sites.

5a (Zone of D. anceps).

The 5a beds of Norway contain a rich coral fauna, especially in
the region of Ringerike, where it has been extensively studied. Since
4ca, the black Tretaspis shale some distance below these 5a beds,
corresponds to the Pleurograptus linearis zone, the 5a beds are
in all probability of the D. anceps zone.

Near Ringerike, at Tyriford, the Rugosa of 5a have been
described by Scheffen (1933). It is characterised by Streptelasmidae
{Streptelasma, Brachyelasma and Grewingkia). The Heliolitidae
however form a better known element of the fauna, thanks to the work
of Kiaer (1899 and 1904) and Lindstrom (1899). In this family there
was an enormous activity in the formation of new genera and species,
many of them with delicate skeletal elements in contrast to the
dominantly coarse skeletal elements of the earliest Heliolitidae. From
Stavnaestangen Trochiscolithus micr aster, Protarea ( Coccoseris )
ungerni, Acidolites asteriscus, Diploepora and Propora bacillifera have
coarse skeletal elements and are associated with the more delicate
Propora affinis, Proheliolites dubius and Plasmoporella convexotabulata
and vesiculosa. At Stord the whole fauna has been listed (Kiaer,
1930). In addition to the Streptelasmidae and Heliolitidae recorded
above, Calapoeciidae occur with Liopora, Calapoecia and Syringo-
phyllum, North American and British genera thus occurring here
together. The Nyctoporinae are represented by Nyctopora cf. billingsi ;
the Halysitidae are present ; the Syringoporidae may well be represented
by Beuschia, and the Favositidae have entered with Palaeofavosites,
with mural pores at the angles, not the middles, of the walls. The
appearance of Palaeofavosites seems the interesting point about this
fauna, which appears a natural development from that of Robeston
Wathen and Coniston, with the American facies represented by
Calapoecia and Nyctopora .

THE ORDOVICIAN CORALS.

The 5b beds of Norway are variously placed, by indirect
correlations, and many writers (Troedsson (1936)) regard them as
basal Llandovery, and separated from 5a by disconformity. Eeliolites
has entered as E. parvistella, and Propora and Proheliolites are the
dominant Heliolitidae. Calapoecia is still present. They will not be
further discussed herein.

Sweden.

Chasmops Limestone.

( f Zone of Climacograptus peltifer or Bier anog rapt us clingani) .

The Chasmops limestone, according to Thorslund (1940) is
developed above the Nemagraptus gracilis zone, its upper part being in
the Dicranograptus clingani zone. The oldest coral in Sweden seems
to be in this limestone, Coelostylis tornquisti Lindstrom (in Angelin
and Lindstrom (1880)), from Dalecarlia and Ostrogothia; from the
longitudinal section it appears to be a columellate streptelasmid, and
its relation to Dalmanophyllum Lang and Smith should be investigated.

Kulsberg Limestone ( ? Zone of B. clingani).

The Kulsberg limestone is below the horizon with Pleurograptus
linearis in the black Trinucleus shales, and is correlated by St^rmer
(1945) with possibly the upper Chasmops limestone, i.e. with the
D. clingani zone. Corals are recorded from the Kulsberg limestone, but
I am unaware of any figures ; it is possibly the lower Leptaena limestone
of Dalecarlia (Thorslund (1935)).

Upper Leptaena Limestone (? Zone of B. anceps).

The upper Leptaena limestone of Dalecarlia is now known as the
Kallholn (= Boda) limestone. The Boda limestone according to
Thorslund (1935) contains a hiatus, the lower part having been
deposited in the Ordovician above the zone of Pleurograptus linearis ,
and the upper part in Silurian times below the zone of Monograptiis
gregarius. Streptelasmids, Propora, Proheliolites, Ealysites, Syringo-
phyllum and Calapoecia are all recorded from it. Thorslund says these
corals seem to come from the upper part of the limestone.

The Southern Lapland Slatdal limestone is coral-bearing and
contains (Kulling (1933)) Rugosa, including a fasciculate form
(“ Columnaria” sp. cf. C. thomi Hall), together with Nyctopora (with
8 long and 8 short septa), Palaeofavo sites, Ealysites, Eofletcheria
and the heliolitids Proheliolites, Propora, Plasmoporella and Eeliolites,
in E. cf. interstincta. Kulling equates this with the Norwegian 5b, the
Balmanites beds of Skarn, the brachiopod beds of Westergotland and
J emtland, the upper Leptaena limestone of Dalarne and the Borkholm
(F2) beds of Estland; and regards it as deposited during an assumed
gap in deposition between the Hartfell and Birkhill of Scotland. It
seems however identical with 'the Norwegian 5a fauna from Stord,
except for the occurrence of Eeliolites, the earliest occurrence of which
elsewhere is in 5b of Norway.

Estland.

C2 ( ? Zone of Climacograptus peltifer).

In the. Kuekers Stage C2 of Estland small solitary streptelasmids
occur, possibly Lamheophyllum with few and rather thin, wavy septa
(S.M.A. 3446). This stage is now correlated with the lower Chasmops
limestone of Sweden and indirectly with the zone of C. peltifer (Thors-
!und 1940),

24 PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

Wesenberg beds ( ? Zone of Pleurograptus linearis).

The Wesenberg beds, which contain the Trentonian Protar aea
vetusta Hall (Lindstrom (1899)) may well be correlated with the Zone
of P. linearis as suggested by Thorslund (1940).

Lyckholm Beds (? Zone of Dicellograptus anceps).

The Lyckholm beds, which are generally correlated with the
Norwegian 5a, contain small Rugosa without dissepiments, Streptelasma,
and “Columnaria” fascicula, with Holy sites, Palaeofavo sites, Syringo-
phyllum, Calapoecia and the heliolitids Trochiscolithus, Protar aea
(Coccoseris) , Propora bacillifera and Propora hirsuta ( = “Nichol-
sonia” megastoma McCoy of Kiaer). This is indeed a 5a fauna (Kiaer
(1899, 1904), Lindstrom (1899)).

The Borkholm beds, F2, which are usually correlated with the
Norwegian 5b, and which it has been suggested are early Silurian,
contain Heliolites parvistella, like the 5b beds, the endemic Palaeopor-
ites, and Propora (Kiaer (1899, 1904), Lindstrom (1899)). They
are not further discussed herein.

Asia.

Spiti (Central Himalayas) Middle Ordovician (Trenton ?).

Reed (1912) regards a fauna from the Pin Valley containing
Streptelasma aff. corniculum, S. sp. and f Liopora sp. as having a
pronounced N. American, flavour and he correlates it with the Mohawk-
ian, suggesting1 Trenton.

Aktscha-tau, Siberia ( ? Caradocian ? Trenton).

Obrutschew* (1926) lists Columnaria alveolata, Ilalysites and
Heliolites interstincta from Aktscha-tau, Siberia, ascribed by Rjabinin
to the Caradocian. If this Heliolites, record is correct this would be the
earliest occurrence of the genius, which otherwise first enters in 5b of
Scandinavia. Lindstrom (1882) lists Calapoecia cribriformis, and
‘Columnaria’ alveolata with a number of Silurian corals from the
Middle Tunguska R., and it seems quite likely that these two names
here apply to Silurian species, though I have seen no figures.

Tasmania.

? D. ancepis zone.

A small fauna from the Queenstown limestone of western Tasmania
(Hill and Edwards (1941)), consisting of a streptelasmid, Tetradium
and the heliolitids Protaraea and Acidolites with a possible Alveolites
sp. is regarded as Upper Ordovician, and shows that corals must have
been able to migrate from the Arctic to the sub-Antarctic regions during
this period. Plasmoporella, Favosites, Favistella and Halysites from
the Chudleigh limestone may also represent an Upper Ordovician fauna,
but a Silurian age is possible for these (Hill (1942) ).

THE ORDOVICIAN CORALS.

IV. REFERENCES.

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Miller, A. K. and Youngquist, W., 1947. Ordovician fossils from the south-western
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Nicholson, H. A., 1875. Report upon the Palaeontology of the Province of Ontario*
96 pp., iv. pis.

1879. On the structure and affinities of the Tabulate Corals of the

Palaeozoic period, xii -j- 342 pp., xv pis. London.

Nicholson, H. A. and Etheridge, R., 1878. A monograph of the Silurian fossils
of the Girvan district in Ayrshire. Fasc. I, pp. 1-135, pis. 1-9.

Nicholson, H. A. and Lydekker, R., 1889. Manual of Palaeontology, 2 Vols., 3rd
Edit., Edinburgh and London.

Oakley, K. P., 1936. An Ordovician species of Chaetetes. Geol. Mag. 73 : 440-444,
pi. xii.

Obrutschew, W. A., 1926. Geologie von Sibirien. Fortschr. Geol. Palaeont. Heft
15, xi + 572, pis.

Okulitch, V. J., 1935. Tetradidae — a revision of the genus Tetradium. Trans, roy.
Soc. Canad. Ill, 29 : Sect. 4, 49-74, pis. i-ii.

1936. Some Chazyan corals. Trans, roy. Soc. Canad. Ill, 30 : Sect.

4, pp. 59-73, pi. 1.

— — ■ 19366. On the genera Heliolites, Tetradium and Chaetetes. Amer.

J. Sci. 32 : 361-379.

1937. Notes on Fletcheria incerta and F. sinclairi. Trans, roy. Canad.

Inst. 21 : 313-6, pi. xviii.

1938. Some Black River Corals. Trans, roy. Soc. Canad. Ill, 32 :

Sect. 4, 87-111, pi. i, ii.

1938. Supposed columella in Tetradium fibratum Safford J. Paleont.

12 : 298.

* 1939. Evolutionary trends of some Ordovician corals. Trans, roy.

, Soc. Canad. Ill, 33 : Sect. 4, 67-80.

1943. The Stony Mountain formation of Manitoba. Trans, roy.

Soc. Canad. (3), 37 : Sect. 4, pp. 59-74, pis. i. ii.

Pringle, J. and George, T. N., 1948. British Regional Geology, South Wales (2nd
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Raymond, P. E., 1905. The fauna of the Chazy Limestone. Amer. J. Sci. 22 : 353-382.

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pp. 72-76.

Reed, F. R. C., 1912. Ordovician and Silurian Fossils from the Central Himalayas.
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Reudeman, R., 1898. Amer. Geol. 22 : 16-25, pi. 5 quoted by Okulitch 1936.

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THE ORDOVICIAN CORALS.

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I, pp. 459-504.

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1938. Geology and palaeontology of the Mingan Islands, Quebec.

Sp. Pap. geol. Soc. Amer. 11 * 132 pp., 24 pis.

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865-888.

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1926. An upper Ordovician fauna from the Rocky Mountains, British

Columbia. Dept. Mines Canad. {Geol. Series No. 46), Bull. No. 44, pp. 1-34.

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during 1925 and 1929. Trans, roy. Soc. Canad. Ill, 20 : Sect. 4, 285-308,
pis. i-v.

Young, F. P., 1943. Black River stratigraphy and faunas. Amer. J. Sci. 241: 141-166,
209-240.

Vol. LXII., No. 2.

APISTOMYIA COLLIN! BEZZI (DIPTERA,
BLEPHAROCERIDAE) IN NORTH
QUEENSLAND.

By I. M. Mackerras and M. J. Mackerras, Queensland Institute of
Medical Research, Brisbane.

( Received 24:th April , 1950; read before the Boy al Society of Queens-
land, 31st July, 1950; issued separately, 5th November, 1951.)

(With Plate I.)

Apistomyia collini Bezzi, 1913, was described from a single female
collected by the late F. P. Dodd at Kuranda, North Queensland, in
September, 1910. So far as we know, it has not been recorded since,
although Mr. D. 0. Atherton has informed us that he has seen Ble-
pharoceridae in North Queensland streams. The following notes are
based on four adults and a considerable series of larvae and pupae, which
we collected near Babinda and Cairns in the same general area as the
type locality. The stages were associated by the method of dissection
described by Tonnoir (1923).

We are indebted to Mr. T. G. Campbell, Division of Economic
Entomology, C.S.I.R.O., Canberra, for the loan of specimens of A.
tonnoiri Tillyard, 1922, for comparison. Specimens of A. collini have
been lodged in the British Museum (Natural History), the Division of
Economic Entomology, C.S.I.R.O., Canberra, and the Institute’s
collection.

Female.

Mr. Paul Freeman of the British Museum (Natural History)
kindly compared one of our specimens with the type in Mr. Collin’s
collection, and reported that “they are indeed the same . . . .” This
sex is to be differentiated most readily from A. tonnoiri by the incon-
spicuous grey dorso-central stripes and barely detectable oblique trans-
verse suture on the scutum, and by the white transverse bands on the
abdomen being complete and not interrupted in the mid line. Bezzi ’s
description and Tonnoir ’s (1930) key are adequate, but the former is
not widely available, so a re-description is given.

Length : Body 5 mm. ; wing 5.5 mm. ; hind leg 13.5 mm.

Head globular. Eyes dark brown, covered with very short, fine,
dense hairs; upper zone of enlarged facets about one-third the depth
of the lower zone. Frons about three-fifths of head width, greyish
black, covered with silvery tomentum, so that the head shows irregular
silvery reflections which vary with the incidence of the light. Ocellar
triangle raised, prominent, velvety black. Antennae a little longer
than the height of the head, ten-segmented, form as in fig. 1 ; basal
segment yellowish to dark brown, silvery at tip, second black, with
silvery apical line, remainder black. Face and basal part of the
elongate proboscis similarly marked to frons, remainder slender, dark
brown ; labella black, long and outwardly coiled ; palpi very short, one-
segmented, black.

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

Thorax. Scutum covered with velvety, jet black to greyish black
tomentum; with narrow grey median and dorso-central lines; and with
the following silvery- white markings : a band across the anterior margin,
conspicuous patches extending obliquely across the anterior half of the
scutum but barely invading the median area between the dorsocentral
lines, joining a silvery- white line just above the lateral margin, and a
silvery triangle in front of the scutellum, which is black basally, silvery
apically (fig. 2) ; the transverse suture is inconspicuous. Pleurae
almost entirely covered with silvery tomentum, but with the sub-alar
area brown and dark brown to blackish patches surrounding the anterior
and posterior spiracles.

Wings only faintly infuscate (not as darkened as would be inferred
from Bezzi’s description), but darker between C and Sc; veins dark
brown to black; the base of Rs is not detectable, An does not reach the
wing margin, and the anal angle is strongly developed, though not as
produced as in the $ of A. mackerrasi Tonnoir, 1930. Halteres with
long yellowish brown stem and large black knob.

Legs with coxae and trochanters yellow; femora brownish yellow
basally, darkening to blackish distally ; remaining segments black. Hind
tibia with a pair of strong apical spurs. Claws long and slender,
finely serrate on basal two-thirds of inner side.

Abdomen. First segment almost entirely covered with silvery
tomentum, but with a narrow black apical transverse patch; second to
fifth covered with velvety black tomentum, and with narrow, complete,
basal, transverse, silvery white bands, which widen considerably at the
sides ; sixth and seventh greyish black, rather shiny, and with relatively
broader silvery bands than on preceding segments; eighth greyish
black; terminal appendages greyish black, obtuse. Venter yellowish
brown.

Male.

Description based on spirit specimens dissected from pupae and
compared with similarly dissected females.

Head large, globular; eyes holoptic, with upper zone of enlarged
facets larger than lower zone ; otherwise similar to $ , except that
mandibles are lacking. Thorax with basic pattern of pigment similar
to $ and differing from dried specimens, as shown in figs. 2, 4, 5. The
silvery tomentum cannot be distinguished in the spirit material of
either sex. Wings and legs crumpled; vein An does not reach the
margin of the wing, and the anal angle appears to be shaped as in the
$ ; spurs are present on the hind tibiae, and the claws bear similar
serrations to the $ . Abdomen with the tergal plates produced
anteriorly to underlie the posterior edge of the preceding segment
(fig. 5) ; whether this indicates that the silvery bands are interrupted
in this sex could not be determined. Hypopygium similar to that
figured by Tonnoir (1930) for A. mackerrasi, and differing from A.
tonnoiri mainly in the relatively shorter, broader ninth tergite and
fused coxites and the proportions of the parts as seen in lateral view
(figs. 6, 7).

The $ of A. collini may be separated from A. mackerrasi by the
incomplete vein An; it is probably best to distinguish it from A.
tonnoiri by the hypopygial characters until fully developed specimens
have been discovered.

APISTOMYIA COLLINI BEZZI.

Pupa.

Length 4 to 4.5 mm. ; breadth 1.7 to 2 mm. ; general form as in fig.
8. Dorsnm dark brown, coarsely tuberculate all over, except on head
■capsule and bases of wing sheaths; indistinct sublateral foveae present
on tergites 3 to 7 and traces on 1 and 2. Outer respiratory lamellae
vertical, broadly triangular, the anterior taller and with rounded apex,
the posterior shorter, truncate. The lateral internal lamella is broadly
triangular, with a rounded or pointed shoulder but no projecting basal
lobe; the medial is acutely triangular (fig. 9). There is some variation
in shape of the lamellae, and the tips appear to have broken off in some
specimens, but all the pupae appear to belong to one species.

The pupa is to be distinguished from that of A. tonnoiri by the
vertical rather than forwardly directed anterior outer respiratory
lamellae, and by the shape of the internal lamellae (cf. our fig. 9 and
fig. 6C of Tonnoir, 1923).

Larva.

Length in last instar 5 to 6 mm. Dorsum dark brown, with
brownish black markings. Antennae black, the distal segment as long
as the basal. Cephalic division finely rugose, with a sublateral pit on
each side in its posterior fifth, a conspicuous black pattern anteriorly
(fig. 11), a row of scale-like spines between the pits, a short black ridge
posterior to these, and a longer black ridge leading to a postero-lateral
row of black spines on each side. Cephalic hairs numerous, projecting
beyond the anterior and antero-lateral margins of the division. The
median divisions have conspicuous black transverse ridges anteriorly
and posteriorly, leading to antero- and postero-lateral rows of strong
black spines. The anterior edge of each division is marked by a row
of broad, brown, scale-like spines, and there are similar but larger
spines between these and the anterior black ridge. The posterior
section of these divisions is unarmed. The anal division has one
transverse black ridge leading to a row of black spines anterior to the
lateral appendage ; a constriction behind the appendage, with a reduced
row of black spines anterior and posterior to it; and a reduced second
appendage, which is dark in colour and bears only two to four long,
strong hairs. There are a few fine submarginal hairs posteriorly on this
division.

The lateral appendages (except the last) are long and conspicuous,
and bear long, dense hairs on their dorsal surface and distal margin ;
the ventral surface and anterior and posterior margins are bare.

Ventral surface creamy brown; suckers normal; ventral gill-tufts
long and slender, three directed forward and two backward; anal gills
globular, the laterals being larger than the medials (fig. 10). Younger
larvae have three gill-filaments (two forward, one backward) and the
cephalic division is relatively larger, but are otherwise similar to the
older ones.

These larvae differ from Tonnoir ’s (1923) figures and description
of A. tonnoiri in the more conspicuous pattern of black pigment, the
larger lateral appendages, bare on their ventral surfaces, and the longer,
slenderer ventral gill-filaments. In some respects they are closer to his
(1930) larva A from Java, but this species has four of the ventral
gill-filaments directed anteriorly, one posteriorly.

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

Habitat.

The early stages were found on rocks and dead timber, where thin
sheets of water were running fast over steep surfaces at the edges of
medium and fairly large cataracts. Some of the larvae in Freshwater
Creek were attached to deeper ledges of rock, where the rush of wafer
was very powerful and the larvae of Gnephia strenua M. & M.
(Simuliidae) occurred in considerable numbers. Four adult females
were captured flying in the spray at the edge of the falls ; no males were
seen. These habitats are typical for the genus, and it is curious that no
specimens have been found so far in apparently equally suitable
situations in southern Queensland.

Distribution.

North Queensland: Kuranda, September, F. P. Dodd (type 2 ) ;

The Cascades, Freshwater Creek, near Cairns, September-October ; The
Boulders, Babinda Creek, near Babinda, September.

The distribution of the genus is linear but curiously discontinuous,
comprising Corsica and Cyprus, the Himalayas, Java, North Queens-
land (the present species), and New South Wales ( A . tonnoiri Till,
from the Blue Mts. and Mt. Kosciusko). Most of the species seem to
be restricted to high country, over 2,000 ft. ; but A. collini is an
exception, in that Kuranda is only 1,080 feet above sea-level and the
other localities are much lower.

Related genera have been found in New Zealand and southern New
South Wales, and the dispersal of the group presents an interesting
problem in zoogeography, on which we do not propose to speculate
here.

REFERENCES.

Bezzi, M., 1913. — Blefaroceridi italiani con descrizione di una nuova forma e di due
specie esotiche. Bull. Soc. ent. Ital., 44: 3-114.

Tillyard, R. J., 1922. — Australian Blepharoceridae. (Order Diptera). Part I: —
Description of new species. Aust. Zool., 2: 159-172.

Tonnoir, A., 1923. — Australian Blepharoceridae. Part II. — Larvae and pupae..
Aust. Zool., 8: 47-59.

Tonnoir, A. L., 1930. — Notes on the genus Apistomyia (Diptera) and description
of a new species. Proc. Linn. Soc. N.S.. Wales, 55: 136-144.

Plate I.

Pkoc. Boy. Soc. Q ’land, Yol. LXII., No. 2.

Apistomyia collini Bezzi. 1, antenna of 9 . 2, dorsal view of thorax of 9 .

3, same of A. tonnoiri for comparison. 4, dorsal view of 9 dissected from pupa.
5, same of ^ . 6, hypopygium of $ , lateral view. 7, same of A. tonnoiri for
comparison. 8, Dorsal view of pupa. 9, respiratory lamellae of pupa (separated
and mounted flat). 10, ventral view of posterior divisions of larva. 11, dorsal
view of larva.

TECHNICAL NOTES.

BANDICOOT FOOD.

Bv Dorothea F. Sahdars, Queensland Institute of Medical Research.

In October 1950, a specimen of Isoodon rnacrourus torosus (Ramsay)
was found dead on a road in Taringa, Brisbane. This was brought to the
Q.I.M.R. Laboratory and subsequently examined for parasites. While
examining the gut, it was noted that much of the contents was still in an
identifiable condition. It consisted mainly of various insects : the muscular
foot region of slugs (these were striped), and a long narrow tail (in 2 pieces)
of a small lizard ( ? Liolepisma) were also recovered.

The insects identified by the Entomology Department of the University
were : — -

Larvae : tenebrionid ; scarabiid ; lepidopteran (2 types) ;

coleopteran (2 types) ; one specimen not identified.

Adult Coleoptera : These were all Harpalinae. By comparison with
specimens in the Queensland Museum they appeared to be : — Gnathophanus
adelaidae Cast., Gnathophanus melanarius Dej., and one specimen which
most closely resembled Hypharpax latiusculus Chaud.

Popular public opinion seems to be that bandicoots burrow for roots
of plants as food.

Most references to the feeding of bandicoots usually describe their
diet as omnivorous. Wood- Jones in “ The Mammals of South Australia ”
says of the bandicoot : — “ They possess an untiring energy in searching
out and destroying insects.” The present case substantiates this. The
statement by Troughton in “ Furred Animals of Australia ” that their
diet is a truly mixed one, of insects, vegetable matter, small mammals and
probably lizards ” is upheld not only by the identification of the contents
of the gut of the animal examined, but also by the fact that the bandicoots
being kept alive at the Q.I.M.R. as laboratory animals, are fed daily on
fresh meat (cut into small pieces) placed on bread. Both are eaten.

The stomach contents of another bandicoot {Isoodon rnacrourus torosus)
collected from Indooroopilly, Brisbane on March 9th, 1951, were mainly
parts of the fruit of P assi flora foetida, the Wild Passion Fruit.

PROCEEDINGS

of the

ROYAL SOCIETY OF QUEENSLAND

VOL. LXII.

C. T. WHITE MEMORIAL SUPPLEMENT

★

The papers that follow are dedicated to the memory of the late
CYRIL TENISON WHITE, Government Botanist of Queensland.

Cyril Tenison White.

I Block by courtesy o f Queensland Naturalists’ Club

Vol. LXIL, No.

CYRIL TEN I SON WHITE, 1890-1950.

Cyril Tenison White was born at Kangaroo Point, Brisbane, on
17th August, 1890, the only child of Mr. and Mrs. Henry White. When
he was five years old the family moved to Rockhampton but later
returned to Brisbane, where he attended the South Brisbane State
School. In 1905 he was appointed Pupil Assistant to his maternal
grandfather, Frederick Manson Bailey, Colonial Botanist of Queens-
land, then in his seventy-eighth year. Some of his early duties were
the copying of letters for his grandfather, the collecting and despatch
of plants of all kinds to overseas specialists and the drawing of nearly
a thousand sketches for Bailey’s Comprehensive Catalogue of Queens-
land Plants. These drawings, crude as they appear to be, give an
unexpectedly accurate impression of the plants and are far more
informative than many illustrations of higher artistic merit. This early
work laid the foundations of White ’s extraordinary knowledge of plants.

F. M. Bailey died in 1915 and was succeeded by his son, John
Frederick Bailey, who had been curator of the Botanic Gardens since
1905. J. F. Bailey held the dual position of Government Botanist and
Curator of the Botanic Gardens until 1917, when he accepted the
position of Director of the Adelaide Botanic Gardens. White then
became Acting Government Botanist, and in 1918 was appointed
Government Botanist, a position he held until his death.

The duties of Government Botanist have always included a large
amount of advisory work for the Department of , Agriculture and Stock
as well as for the general public, and White’s knowledge of cultivated
plants was freely called upon. At first White carried on almost alone,
but in 1919 W. D. Francis was appointed assistant, and it was not
until the 1930 ’s that other botanists joined his staff. He was Lecturer
in Forest Botany in the University of Queensland and had been a
member of its Faculty of Agriculture.

White had a wide interest in science, horticulture, and the conserva-
tion of nature, and took an active part in many societies. He served
on the council of the Royal Society of Queensland from 1909 until
1911 and, except for occasional years, from 1918 to 1938; he was
honorary librarian in 1909-11 and president in 1921. He took a special
interest in the Queensland Naturalists’ Club and he served on its
council from 1910 until his death, except for the single year 1939,
when he was in England ; he was honorary excursion secretary in 1912,
honorary secretary and treasurer in 1913, honorary secretary in 1914-7,
honorary editor from 1920 to 1938 and in 1940 (sharing the position
with A. H. Chisholm in 1920-22), and president in 1918, 1923, and
1943. He was a regular attendant at excursions (which he often led)
and meetings, and stimulated interest in young and old alike. From
1922 to 1927 he was local honorary secretary to the Australasian
Association for the Advancement of Science, and he served on the
councils of the Queensland Horticultural Society, Royal Geographical
Society of Australasia (Queensland Branch), Queensland Orchid
Society, Professional Officers’ Association (of all of which he had been
president), and the National Parks Association of Queensland. He
had been a Fellow of the Linnaean Society of London, but resigned

36 PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

his membership some years ago. He was awarded the Mueller Medal
for distinguished services to Australian Natural History in 1946 and
was granted the honorary degree of Master of Science by the University
of Queensland in 1948.

White loved plants and had an amazing knowledge of the flora
of Australia, New Caledonia, and New Guinea, and of garden plants.
He made many private collecting trips in unofficial time. He collected
in all Australian States except the Northern Territory, in Papua (in
1918, chiefly in the neighbourhood of Yule Island, Port Moresby and
the Astrolabe Range), in New Caledonia in 1923, England in 1939,
the Territory of New Guinea in 1944 when he acted as Instructor to
Army Forestry Units, and in 1945, and he assisted in a forestry survey
of the Solomon Islands in 1945. His numbered collections exceed
13,000, but the specimens collected before his Papuan trip of 1918
were not numbered. The collections in the Territory of New Guinea
carry mostly N.G.F. (New Guinea Forests) numbers and the collections
made with F. S. Walker in the Solomon Islands form a series preceded
by the letters B.S.I.P. Some small collections made in Queensland
also have different numberings. His collections greatly expanded the
Queensland Herbarium started by F. M. Bailey, and his duplicates are
widely distributed. Early in 1939 he went to England as Australian
Liaison Officer at the Royal Botanic Gardens, Kew, but war conditions
forced him to return to Brisbane before the end of the year.

In spite of the calls made on his time by an ever-increasing amount
of routine matters, White made a very large contribution to the
knowledge of the floras of Queensland, north-eastern New South
Wales, New Caledonia, New Guinea, and the Solomon Islands. He had
an extensive knowledge of all groups of vascular plants, but he was
particularly interested in woody plants. Some of his papers suggest
that he was chiefly interested in the discovery and recording of what
was new or unusual ; this may have been the case in his younger days,
but a great part of his published reports of plants new to Queensland
or of new species was based on fairly extensive studies of all available
material of the genera concerned. He worked rapidly but carefully
and was always eager to send material to specialists for detailed study.

His earliest publications were in the Queensland Naturalist , and
to this journal he made nearly fifty contributions, chiefly brief accounts
of the vegetation of localities visited or plants collected during the
excursions of the Queensland Naturalists’ Club, but also an illustrated
account of twenty-three species of Eucalyptus found in the neighbour-
hood of Brisbane and keys to twenty species of Acacia from the same
area. By far the greatest number of articles with which his name is
associated appeared in the Queensland Agricultural Journal. These
are mostly short and often semipopular in nature; nearly a hundred
deal with weeds or plants suspected poisonous to stock or humans,
and a few of these were written with J. F. Bailey, F. Smith, or E. H.
Gurney; forty others, with W. D. Francis as co-author, are well-
illustrated accounts of Queensland trees that were afterwards used
by Francis in his Australian Bain-forest Trees. The bulk of his
systematic work on Queensland plants appeared under the title of
“Contributions to the Queensland Flora”; the earlier ones, with
J. F. Bailey or W. D. Francis as co-author, appeared as Botany
Bulletins of the Queensland Department of Agriculture and Stock,

CYRIL TENISON WHITE.

but from 1922 onwards they appeared in the Proceedings of the Royal
Society of Queensland , with W. D. Francis as co-author of the earlier
of these. Another large paper is “ Ligneous plants collected for the
Arnold Arboretum in North Queensland by S. F. Kajewski in 1929/’
published as Contributions from the Arnold Arboretum No. 4. Most
of his earlier work on the flora of New Guinea was published in the
Proceedings of the Royal Society of Queensland (that on the collections
of Lane-Poole with W. D. Francis), but his later work on this area,
on New Caledonia, and the Solomon Islands appeared in the Journal
of the Arnold Arboretum. (A preliminary account of his collections
from the Solomon Islands was given by F. S. Walker in his report:
“The Forests of the British Solomon Islands Protectorate”; London,
1948.) Three biographical papers give an interesting account of early
Queensland botany and incidentally of his own background. The
“Tryon Memorial Lecture,” in 1945, dealt with H. Tryon, the others,
in 1945 and 1950, dealt with F. M. Bailey and his family.

White wrote two useful books. 1 ‘ An elementary textbook of
Australian forest botany ’ ’ was published in 1922, and a second edition
appeared in 1925. Both editions are marked “Vol. 1”; a second volume
was planned but not written. “Principles of Botany for Queensland
Farmers” was published in book form in 1941 but had previously
appeared in seven parts in the Queensland Agricultural Journal in
1936-37. Both are textbooks for botany in their respective fields, with
Australian examples, and both give good general accounts of Queensland
vegetation.

His kindliness, tolerance, good humour, and enthusiasm endeared
“C.T.” to a large number of friends and acquaintances from many
walks of life. He was always ready and willing to give advice, but
rarely offered if unless asked; he never forced his opinions on anybody
and his kindliness was often imposed on. He was a skilful raconteur,
and his wealth of anecdote enlivened many a lunch period in the
herbarium and camp-fire gathering. He was a link between the old
school of naturalists and the modern one trending to specialization
and was able to adjust himself to the change.

Almost his only hobbies were his home and his garden, and his
wife and two daughters shared his interest in natural history. He
was seriously ill for some months in 1947, but recovered sufficiently
to carry on most of his usual activities at a reduced tempo. A heart
attack on the eve of his sixtieth birthday meant the loss of a lovable
man and an outstanding scientist who, on the day of his death, was
preparing for a collecting trip to North Queensland.

The following bibliography is based on a list of publications
prepared by Mr. J. Clancy. In addition to the works listed, notes
on a variety of plants appeared in many parts of the Queensland
Agricultural Journal in. 1926-46 (new series, vols. 26-63) under the
heading “Answers to Correspondents.” White’s name has been also
associated elsewhere in the same journal with the identification of plants
and remarks thereon.

S. T. BLAKE,

Botanic Museum and Herbarium,

Botanic Gardens, Brisbane.

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

BIBLIOGRAPHY.

1908.

List of Plants secured during Sankey’s Scrub Excursion. Queensl. Nat. 1 : 56.

Excursion to Bulwer, Moreton Island, 12th- 14th September, 1908. — Botany. QueensL
Nat. 1 : 70-73. (With J. Wedd.)

1910.

List of Plants collected at Glasshouse Mountains, September 11th to 13th, 1909.
Queensl. Nat. 1 : 119-120. (With J. Wedd.)

List of Plants collected in addition to those obtained in September, 1909. Queensl.
Nat. 1 : 154-155. (With J. Wedd.)

1911.

Botanic Notes, No. 1. Queensl. Nat. 1 : 204-208.

1913.

Botanic Notes, No. 2. Queensl. Nat. 1 : 256-261.

1914.

Eleusine indica (crowfoot grass) — A grass dangerous to stock. Queensl. Agric. J. n.s. 2 t
158-160. (With F. Smith.)

1915.

Notes on a few interesting plants from Moreton Bay. Proc. Roy. Soc. Queensl. 21 :
96-99.

Botanic Notes, No. 3. Queensl. Nat. 2 : 21-23.

Illustrated notes on the weeds of Queensland, No. 1. On three closely allied weeds—
Aster subulatus Michx., Erigeron canadensis Linn., and Erigeron linifolius
Willd. Queensl. Agric. J. n.s. 4 : 226-230. (With J. F. Bailey.)

Plants poisonous to stock. Dysphania myriocephala Benth. (N.O. Illecebraceae).
Queensl. Agric. J. n.s. 3 : 264-265. (With F. Smith.)

Contributions to the flora of Queensland. New Series, No. 1. Queensl. Agric. J. n.s. 4 :
287-289. (With J. F. Bailey.)

1916.

Notes on a few interesting plants collected in the vicinity of Brisbane. Proc. Roy.
Soc. Queensl. 28 : 107-111.

Contributions to the Queensland Flora. Queensl. Dep. Agric. Bot. Bull. 18. (With
J. F. Bailey.) 16 pp.

Illustrated notes on the weeds of Queensland, No. 2. Cassia occidentals Linn, (order
Leguminosae). Queensl. Agric. J. n.s. 5 : 40-42. (With J. F. Bailey.)

Illustrated notes oil the weeds of Queensland. No. 3. Khaki Weed ( Alternanthera
achyrantha R.Br.) Queensl. Agric. J. n.s. 5 : 277-278. (With J. F. Bailey.)

Illustrated notes on the weeds of Queensland. No. 4. On two species of Gomphocarpus
(Wild Cotton) naturalized in Queensland. Queensl. Agric. J. n.s. 6 : 104-106.
(With J. F. Bailey.)

Illustrated notes on the weeds of Queensland. No. 5. Sida acuta Burm. (Order
Malvaceae). Queensl. Agric. J. n.s. 8 : 262-263. (With J. F. Bailey.)

Illustrated notes on the weeds of Queensland. No. 6. Cape Spinach, Emex australis
Steinh. Queensl. Agric. J. n.s. 6 : 373-374. (With J. F. Bailey.)

1917.

A new species of Xanthostemon (nat. ord. Myrtaceae) from northern Queensland.
Proc. Roy. Soc. Queensl. 29 : 57-59.

The flora of a single tree. Proc. Roy. Soc. Queensl. 29 : 64-69. (With H. A. Longman.)
Botanic notes, No. 4. Queensl. Nat. 2 : 65-66.

Brief report on the botany of the excursion to Buderim Mountain, Easter, 1916.
Queensl. Nat. 2 : 67.

Report on the botany of excursion to My ora, Stradbroke Island, Easter, 1917. Queensl.
Nat. 2 : 67-68.

Illustrated notes on the weeds of Queensland. No. 7. “ Grass Seed ” or “ Mackie’s

Pest ” (Chrysopogon aciculatus Trin.). Queensl. Agric. J. n.s. 1 : 246-247.

CYRIL TENISON WHITE.

Illustrated notes on the weeds of Queensland. No. 8. “ Giant pig Weed ” ( Trianthema

portulacastrum Linn.). Queensl. Agric. J. n.s. 7 : 290-291.

Illustrated notes on the weeds of Queensland. No. 9. On the species of Datura (Thorn
Apple) naturalised in Queensland. Queensl. Agric. J. n.s. 8 : 31-35.

Illustrated notes on the weeds of Queensland. No. 10. Sida cordifolia Linn. Queensl.
Agric. J. n.s. 8 : 84-85.

Illustrated notes on the weeds of Queensland. No. 11. “ Devil’s Fig ” ( Solatium

largiflorum, n.sp.). Queensl. Agric. J. n.s. 8 : 170-172.

Plants poisonous to stock. Tape Vine ( Stephania hernandiaefolia Walp.). Queensl.
Agric. J. n.s. 8 : 230-232.

Records of a few alien plants. Queensl. Agric. J. n.s. 8 : 269-270.

Contributions to the Queensland Flora. Queensl. Dep. Agric. Bot. Bull. 19. (With
J. F. Bailey.) 20 pp.

1918.

An interim census of cyanophoric plants in the Queensland flora. Proc. Roy. Soc.
Queensl. 30 : 84-90. (With F. Smith.)

Mutation in a proteaceous tree. Proc. Roy. Soc. Queensl. 30 : 162-165. (With H. A.
Longman.)

Contributions to the Queensland flora. Queensl. Dep. Agric. Bot. Bull. 20. (20 pp.)

Illustrated notes on the weeds of Queensland. No. 12. Twiggy Mullein ( Verbascum
virgatum With.). Queensl. Agric. J. n.s. 9 : 22-23.

Illustrated notes on the weeds of Queensland. No. 13. Mossman River Grass ( Cenchrus
echinatus Linn.). Queensl. Agric. J. n.s. 9 : 180-181.

Weeds and poisonous plants of the Atherton Tableland. Queensl. Agric. J. n.s. 9 :
147-155.

Records of a few alien plants. Queensl. Agric. J. n.s. 9 : 228.

On a peculiar subterranean fruiting habit of Vigna lanceolata R.Br., with description
of a new variety. Queensl. Agric. J. n.s. 10 : 41-44.

Two native leguminous fodder plants. Queensl. Agric. J. n.s. 10 : 252-254.

1919.

On the occurrence of abortive styles in Buckinghamia celsissima F.v.M. Proc. Roy.
Soc. Queensl. 31 : 42-45. (With C. D. Gillies.)

A revised account of the Queensland Lecythidaceae. Proc. Linn. Soc. N. S. Wales 44 :
822-825.

Illustrated notes on the weeds of Queensland. No. 14. Prickly poppy ( Argemone
mexicana Linn. var. ochroleuca Lindl.). Queensl. Agric. J. n.s. 11 : 118-119.

Illustrated notes on the weeds of Queensland. No. 15. On two species of Labiatae
naturalised in northern Queensland. Queensl. Agric. J. n.s. 12 : 141-143.

Illustrated notes on the weeds of Queensland. No. 16. A “ rattlepod ” ( Crotolaria
sericea Retz.). Queensl. Agric. J. n.s. 12 : 198-199.

Luminous fungi. Queensl. Agric. J. n.s. 12 : 33-34.

Lomatia silaifolia — A poisonous flower. Queensl. Agric. J. n.s. 11 : 256-257.

Miscellaneous botanical notes. (A) Notes on a few native plants. (B) Records of a
few alien plants. Queensl. Agric. J. n.s. 12 : 75.

Records of poisoning of birds by two species of Cassia. Queensl. Agric. J. n.s. 12 : 306.
Contributions to the Queensland flora. Queensl. Dep. Agric. Bot. Bull. 21 : (23 pp.)

1920.

On the occurrence of cyanophoric glucosides in the flowers of some Proteaceae. Proc.
Roy. Soc. Queensl. 32 : 89-91. (With F. Smith.)

Contributions to the orchidaceous flora of Queensland. Proc. Roy. Soc. Queensl. 32 :
117-124. (With R. S. Rogers.)

The peach-leaf poison bush, Trema aspera Blume : Its occasional toxicity. Proc. Roy.
Soc. Queensl. 32 : 132-134. (With F. Smith.)

Two interesting fungi. Queensl. Nat. 2 : 80-82.

Natural History Exhibition — Botany. Queensl. Nat. 2 : 90-92.

A rare and beautiful native Tecoma. Queensl. Nat. 2 : 99-102.

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

Flora of the Bunya Mountains. Queensl. Agric. J. n.s. 13 : 25-31.

Botanical notes. (A) Description of a new variety of the red cedar. (B) On a Haloragis
not previously recorded from Queensland. Queensl. Agric. J. n.s. 13 : 66-67.

Two native drought-resistant fodder plants. Queensl. Agric. J. n.s. 13 : 172-175.

Results of feeding experiments with a suspected poisonous plant ( Wikstroemia indica
C. A. Mey.). Queensl. Agric. J. n.s. 13 : 172-175. (With C. J. Pound.)

The red ash ( Alphitonia excelsa), a valuable fodder tree. Queensl. Agric. J. n.s. 13 :
218-219.

The weir vine (Ipomoea calobra). With notes on its reputed poisonous properties, based
on information supplied by Donald Gunn, Esq., M.L.A. Queensl. Agric.
J. n.s. 13 : 269-272.

Note on variation in the bark of two common eucalypts. Queensl. Agric. J. n.s. 14 :
70-72.

The White Cedar ( Melia azedarach var. australasica ) : A plant poisonous to pigs.
Queensl. Agric. J. n.s. 14 : 146-147.

Illustrated notes on the weeds of Queensland. No. 18. Onion weed ( Asphodelus
fistulosus Linn.). Queensl. Agric. J. n.s. 14 : 196-197.

Illustrated notes on the weeds of Queensland. No. 19. Burr buttercup ( Ranunculus
muricatus L.). Queensl. Agric. J. n.s. 14 : 243-244.

On two species of kurrajong ( Brachychiton ) occurring in Queensland. Queensl. Agric.
J. n.s. 14 : 289-293.

Contributions to the Queensland Flora. Queensl. Dep. Agric. Bot. Bull. 22. (With
W. D. Francis.) 36 pp.

1921.

Contributions to the orchidaceous flora of Queensland, No. 2. Proc. Roy. Soc. Queensl.
32 : 135-143. (With R. S. Rogers).

Contributions to the Queensland flora. Proc. Roy. Soc. Queensl. 33 : 152-165. (With
W. D. Francis.)

Botany of Stradbroke Island. Queensl. Nat. 2 : 124-126.

Four notable native plants. Queensl. Nat. 3 : 14-17.

Notes on mistletoes. Queensl. Nat. 3 : 39.

Three interesting fungi. Queensl. Nat. 3 : 37-38.

The flora of a small tropical island by W. Bradke : Foreword by C. T. White. Queensl.
Nat. 3 : 133-134.

Notes on the genus Flindersia (family Rutaceae). Proc. Linn. Soc. N. S. Wales 46 :
324-329.

The pongamia tree ( Pongamia glabra)- — A useful fodder tree. Queensl. Agric. J. n.s.
15: 27-28.

A wild cotton. Queensl. Agric. J. n.s. 15 : 216-218.

The carob and algaroba beans. Queensl. Agric. J. n.s. 15 : 266-269.

Illustrated notes on the weeds of Queensland. No. 20. Star burr ( Acanthospermum
hispidum DC.). Queensl. Agric. J. n.s. 15 : 126-127.

Illustrated notes on the weeds of Queensland. No. 21. Goosefoot ( Chenopodium
triangulare R.Br.). Queensl. Agric. J. n.s. 15 : 171-172.

Illustrated notes on the weeds of Queensland. No. 22. Indian heliotrope ( Heliotropium
indicum Linn.). Queensl. Agric. J. n.s. 16 : 103-104.

Illustrated notes on the weeds of Queensland. No. 23. Snake weed (Stachytarpheta
dichotoma). Queensl. Agric. J. n.s. 16 : 194-195.

Illustrated notes on the weeds of Queensland. No. 24. Wild sage ( Salvia verbenaca) .
Queensl. Agric. J. n.s. 16 : 267-268.

Two plants poisonous to stock. Queensl. Agric. J. n.s. 16 : 194-196.

A native yam. Queensl. Agric. J. n.s. 16 : 386.

Queensland trees. No. 1. Scrub ironbark ( BHdelia exaltata). Queensl. Agric. J. n.s.
15 : 173-175. (With W. D. Francis.)

Queensland trees. No. 2. Giant water gum ( Eugenia francisii). Queensl. Agric.
J. n.s. 15 : 219-221. (With W. D. Francis.)

CYRIL TENISON WHITE.

Queensland trees. No. 3. Lignum-vitae ( Vitex lignum-vitae). Queensl. Agrie. J. n.s.

15 : 270-272. (With W. D. Francis.)

Queensland trees. No. 4. Yellow sassafras (Doryphora sassafras). Queensl. Agrie.
J. n.s. 16 : 43-45. (With W. D. Francis.)

Queensland trees. No. 5. Blue berry ash ( Elaeocarpus obovatus). Queensl. Agrie.
J. n.s. 16 : 105-107. (W. D. Francis.)

Queensland trees. No. 6. The coondoo ( Sideroxylon Richardi). Queensl. Agrie. J. n.s.

16 : 197-199. (With W. D. Francis.)

Queensland trees. No. 7. Koda ( Ehretia acuminata). Queensl. Agrie. J. n.s. 16 :
271-273. (With W. D. Francis.)

1922.

A contribution to our knowledge of the flora of Papua (British New Guinea). Proc.
Roy. Soc. Queensl. 34 : 5-65. (Presidential Address.)

An elementary textbook of Australian forest botany, vol. 1. Govt. Printer, Sydney.
223 pp., 105 figs.

Illustrated notes on the weeds of Queensland. No. 25. A western burr ( Sida platy calyx) .
Queensl. Agrie. J. n.s. 17 : 34-35.

Illustrated notes on the weeds of Queensland. No. 26. Wild salvia ( Salvia coccinea).
Queensl. Agrie. J. n.s. 17 : 70-71.

Illustrated notes on the weeds of Queensland. No. 27. Tumbling mustard or oriental
rocket {Sisymbrium orientate). Queensl. Agrie. J. n.s. 18 : 362-363.

Weeds of Queensland. No. 28. The jo-jo weed ( Soliva sessilis), a new lawn pest.

Queensl. Agrie. J. n.s. 18 : 398-399.

Zamia — Comment. Queensl. Agrie. J. n.s. 7 : 53.

The Algaroba bean ( Prosopis juliflora) : Is it likely to become a pest — Comment.
Queensl. Agrie. J. n.s. 17 : 271-272. (With others.)

A mangosteen from north Queensland. Queensl. Agrie. J. n.s. 18 : 34-35.

A native fodder tree from north Queensland. Queensl. Agrie. J. n.s. 18 : 96-97.

Queensland trees. No. 8. Opossum wood ( Quintinia sieberi). Queensl. Agrie. J. n.s.

17 : 72-74. (With W. D. Francis.)

Queensland trees. No. 9. White myrtle ( Rhodamnia argentea). Queensl. Agrie. J. n.s.
17 : 107-109. (With W. D. Francis.)

Queensland trees. No. 10. Socket wood (Daphnandra micrantha). Queensl. Agrie.
J. n.s. 17 : 230-232. (With W. D. Francis.)

Queensland trees. No. 11. Yellow- wood {Flindersia Oxleyana). Queensl. Agrie. J. n.s.
17 : 304-306. (With W. D. Francis.)

Queensland trees. No. 12. The ivorywood. Queensl. Agrie. J. n.s. 18 : 45-47. (With
W. D. Francis.)

Queensland trees. No. 13. The crow’s apple. Queensl. Agrie. J. n.s. 18 : 89-91. (With
W. D. Francis.)

Queensland trees. No. 14. Pennantia cunninghamii. Queensl. Agrie. J. n.s. 18 :
159-161. (With W. D. Francis.)

Queensland trees. No. 15. The hauer {Dissiliaria baloghioides) . Queensl. Agrie.
J. n.s. 18 : 359-361. (With W. D. Francis.)

Queensland trees. No. 16. Red Carabeen. Queensl. Agrie. J. n.s. 18 : 394-396. (With
W. D. Francis.)

1923.

Contributions to the Queensland Flora. No. 2. Proc. Roy. Soc. Queensl. 35 : 63-84.
(With W. D. Francis.)

A reputed blue-flowering variety of Portulaca bicolor. Queensl. Nat. 4 : 40.

A new conifer from southern Queensland. Proc. Linn. Soc. N. S. Wales 48 : 449-450.

Weeds of Queensland. No. 29. The peach-leaf poison bush or wild peach ( Trema
aspera). Queensl. Agrie. J. n.s. 19 : 81-82.

Weeds of Queensland. No. 29 (bis). Solanum seaforthianum. Queensl. Agrie. J. n.s.
19 : 238-239.

Weeds of Queensland. No. 31. The rubber vine ( Cryptostegia grandiflora) . Queensl.
Agrie. J. n.s. 19 : 286-287.

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

Weeds of Queensland. No. 32. Cleome aculeata. Queensl. Agric. J. n.s. 19 : 516-517.

Weeds of Queensland. No. 33. Tree groundsel ( Baccharis Tialimifolia) . Queensl.
Agric. J. n.s. 20 : 49-50.

Weeds of Queensland. No. 34. A “ rattle pod ” Crotolaria striata. Queensl. Agric.
J. n.s. 20 : 236-239.

Sour grass or yellow grass. Queensl. Agric. J. n.s. 20 : 46.

A thin-shelled variety of the Queensland nut ( Macadamia ternifolia). Queensl. Agric.
J. n.s. 20 : 93-95.

Queensland trees. No. 17. The black apple or black plum ( Sideroxylon australe ).
Queensl. Agric. J. n.s. 19 : 78-80. (With W. D. Francis.)

Queensland trees. [No. 18.] Southern ghittoe ( Halfordia drupifera). Queensl. Agric.
J. n.s. 19 : 197-199. (With W. D. Francis.)

Queensland trees. No. 19. The scrub beefwood. Queensl. Agric. J. n.s. 19 : 304-305,
415. (With W. D. Francis.)

Queensland trees. No. 20. [The rose-leaf or pink marara, AcJcama paniculata.] Queensl.
Agric. J. n.s. 19 : 409-441. (With W. D. Francis.)

Queensland trees. No. 21. The maiden’s blush. Queensl. Agric. J. n.s. 19: 513-515.
(With W. D. Francis.)

Queensland trees. No. 22. [The bolly gum, Litsea reticulata.~\ Queensl. Agric. J. n.s.
20 : 46-48. (With W. D. Francis.)

Queensland trees. No. 23. The corduroy tamarind. Queensl. Agric. J. n.s. 20 : 90-92,
(With W. D. Francis.)

Queensland trees. No. 24. The blue quandong. Queensl. Agric. J. n.s. 20 : 175-177.
(With W. D. Francis.)

Queensland trees. No. 25. Pepperberry tree. Queensl. Agric. J. n.s. 20 : 362-364.
(With W. D. Francis.)

Queensland trees. [No. 26. Ribbonwood, Euroschinus falcatus.'] Queensl. Agric. J. n.s.

20 : 434-436. (With W. D. Francis.)

1924.

On a new species of Melaleuca (family Myrtaceae) from southern Queensland. Proc.
Roy. Soc. Queensl. 36 : 41-43. (With E. Cheel.)

The eucalypts or gum trees of the Brisbane district. Queensl. Nat. 4 : 67-72.

The eucalypts or gum trees of the Brisbane district, II. Queensl. Nat. 4 : 109-112.

Flora of Moreton Island. (Report on the Easter encampment (1924) of the Queensland
Naturalists’ Club at Cowan Cowan.) Queensl. Nat. 4 . 86-91.

Weeds of Queensland. No. 35. Bitter bark, native cinchona or quinine ( Alstonia
constricta). Queensl. Agric. J. n.s. 21 : 252-253.

Weeds of Queensland. No. 36. Bluetop heliotrope (Heliotr opium anchusaefolium) .
Queensl. Agric. J. n.s. 23. : 284-285.

Notes on two weed pests. Queensl. Agric. J. n.s. 21 : 286.

Wattle bark : The possibilities of its cultivation in Queensland. Queensl. Agric. J. n.s.
21: 423-425.

The southern olive, Notelaea longifolia var. velutina. A native plant harbouring fruit-fly
larvae in the Stanthorpe district. Queensl. Agric. J. n.s. 22 : 240-241.

Possibilities of camphor cultivation in Queensland. Queensl. Agric. J. n.s. 22 : 308.
Two valuable fodder trees. Queensl. Agric. J. n.s. 22 : 352-356.

The native bryony ( Bryonia laciniosa ). A poisonous plant. Queensl. Agric. J. n.s. 22 :
442-444.

Queensland trees. [No. 28. South Queensland kauri, Agathis robusta ] Queensl.
Agric. J. n.s. 21 : 34-36. (With W. D. Francis.)

Queensland trees. [No. 29. Sour cherry, Eugenia corynantha .] Queensl. Agric. J. n.s.

21 : 281-283. (With W. D. Francis.)

Queensland trees. No. 30. [ Cryptocarya patentinervis.\ Queensl. Agric. J. n.s. 21 :

437-439. (With W. D. Francis.)

Queensland trees. No. 31. [Bunya pine, Araucaria Bidwillii ; a bolly gum,
Beilschmiedia elliptical Queensl. Agric. J. n.s. 22 : 26-27, 32. (With W. D.
Francis.)

CYRIL TENISON WHITE.

Queensland trees. [No. 32. Sideroxylon pohlmanianum . ] Queensl. Agric. «J. n.s. 22 :
G5-67. (With W. D. Francis.)

Queensland trees. [No. 33. Lucuma amorphosperina.] Queensl. Agric. J. n.s. 22 :
238-239. (With W. D. Francis.)

Queensland trees. [No. 34. Endiandra compressa.] Queensl. Agric. J. n.s. 22 : 308-310.
(With W. D. Francis.)

Queensland trees. [No. 35. Broad-leaved leopard tree, Flindersia collina .] Queensl.
Agric. J. n.s. 22 : 356-358. (With W. D. Francis.)

Queensland trees. [No. 36. Native tamarind, Diploglottis ciinninghamii.] Queensl.
Agric. J. n.s. 22 : 444-446. (With W. D. Francis.)

1925.

An elementary textbook of Australian forest botany, vol. 1. 2nd ed. Govt. Printer,
Sydney.

The eucalypts or gum trees of the Brisbane district, III. Queensl. Nat. 5 : 12-16.

The eucalypts or gum trees of the Brisbane district, IV. Queensl. Nat. 5 : 43-45.

Fruit of Acronychia imperforata — A “ wild orange.” [Remarks on a note by J. H.
Schmeider.] Queensl. Nat. 5 : 48.

Weeds of Queensland. No. 37. Devil’s claw ( Martynia lutea). Queensland Agric.
J. n.s. 23 : 96-97.

Weeds of Queensland. No. 38. Bassia birchii (a “ Chinese burr ”). Queensl. Agric.
J. n.s. 23 : 200-201.

Weeds of Queensland. No. 39. Narrow-leaved sage ( Salvia lancifolia). Queensl.
Agric. J. n.s. 23 : 417-418.

Weeds of Queensland. No. 40. American dog weed ( V erbescinci encelioides). Queensl.
Agric. J. n.s. 23 : 520-521.

Vinca rosea — A reputed cure for diabetes. Queensl. Agric. J. n.s. 23 : 143-144.
Queensland forests and forest trees. Queensl. Agric. J. n.s. 24 : 124-128.

Queensland trees. [No. 37. Scrub wilga, Geijera salicifolia .] Queensl. Agric. J. n.s. 23 :
146-147. (With W. D. Francis.)

Queensland trees. [No. 38. Deep yellow-wood, Rhodosphaera rhodanthema.] Queensl.
Agric. J. n.s. 23 : 202-204. (With W. D. Francis.)

Queensland trees. [No. 39. Cryptocarya foveolata.] Queensl. Agric. J. n.s. 23 : 334-335.
(With W. D. Francis.)

Queensland trees. [No. 40. Australian olive, Olea paniculata .] Queensl. Agric. J. n.s,
24 : 438-440. (With W. D. Francis.)

1926.

Contributions to the Queensland flora, No. 3. Proc. Roy. Soc. Queensl. 37 : 152-167,
(With W. D. Francis.)

The eucalypts or gum trees of the Brisbane district, V. Queensl. Nat. 5 : 51-53.
The eucalypts or gum trees of the Brisbane district, VI. Queensl. Nat. 5 : 94-96.

A previously undescribed Dodonaea from ‘south-eastern Queensland. Queensl. Nat. 6 :
13-14.

Report on the botany of the Queensland Naturalists’ Club Excursion to Elimbah,
11th- 13th Sept., 1926. Queensl. Nat. 6 : 14-15.

On a small collection of plants from the Rigo district, Papua (British New Guinea).
Proc. Linn. Soc. N. S. Wales 51 : 296-298.

A new species of Diplospora from southern Queensland. J. Bot. 64 : 216-217.

The genus Sonneratia in Queensland. J. Bot. 64 : 217-218.

A variety of Ceriops tagal C. B. Rob. ( C . candolleana W. & A.). J. Bot. 64 : 220-221.

Ligneous plants collected in New Caledonia by C. T. White in 1923. .T. Arnold Arb. 7 :
74-103. (With E. H. Wilson and A. Guillaumin.)

A new species of Paramignya of Papua with notes on two other Papuan Rutaceae,
J. Arnold Arb. 7': 231-233.

An interesting plant, Gomphrena leontopodioides . Queensl. Agric. J. n.s. 25 : 270 271.
Destruction of the khaki weed. Queensl. Agric. J. n.s. 25 : 512,

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

Weedfi of Queensland. No. 41. Rivina (Rivina laevis). Queensl. Agric. J. n.s. 25 :
274-275.

Weeds of Queensland. No. 42. Milky cotton bush ( Asclepias curassavica). Queensl.
Agric. J. n.s. 26 : 125-127.

1927.

Plants collected in Papua by C. E. Lane-Poole. Proc. Roy. Soc. Queensl. 38 : 225-201.
(With W. D. Francis.)

A previously undescribed species of Maniltoa from Papua. J. Arnold Arb. 8 : 130.

Two Papuan species of Ardisia. J. Bot. 65 : 248.

Rivina ( Rivina laevis). Queensl. Agric. ,T. n.s. 27 : 130.

Vinca rosea — A reputed cure for diabetes. Queensl. Agric. J. n.s. 28 : 354-355.

Queensland trees. [No. 41.] The tallow-wood ( Eucalyptus microcorys). Queensl.
Agric. J. n.s. 28 : 46-48.

1928.

Plants collected in the Mandated Territory of New Guinea by C. E. Lane-Poole. Proc.
Roy. Soc. Queensl. 39 : 61-70. (With W. D. Francis.)

The eucalypts or gum trees of the Brisbane district, VII. Queensl. Nat. 6 : 78-82.
Recent botanical work in the Pacific. Queensl. Nat. 6 : 85-88.

Weeds of Queensland. Button weed or button mallow ( Modiola caroliniana) . Queensl.
Agric. J. n.s. 30 : 598-599.

1929.

Contributions to the Queensland flora, No. 4. Proc. Roy. Soc. Queensl. 41 : 139-143.
(With W. D. Francis.)

The eucalypts or gum trees of the Brisbane district, VIII. Queensl. Nat. 7 : 36-39.

Ligneous plants collected in the Territory of Papua (British New Guinea) in 1925-26
by L. J. Brass. J. Arnold Arb. 10 : 197-274.

William Vincent Fitzgerald. J. Bot. 67 : 309.

Weeds of Queensland. The correct botanical identity of the Lantana naturalised in
Queensland. Queensl. Agric. J. n.s. 31 : 294-296.

Weeds of Queensland. Cassia laevigata Willd. (an arsenic bush). Queensl. Agric. J. n.s.
31 : 352-353.

Queensland weeds. Twin leaf ( Zygophyllum apiculatum) ; Solanum auriculatum—a
“ wild tobacco.” Queensl. Agric. J. n.s. 32 : 194-197.

1930.

Two interesting Queensland eucalypts. Proc. Roy. Soc. Queensl. 42 : 82-85. (With ,
W. F. Blakely).

The margosa tree and its allies. Queensl. Agric. J. n.s. 34 : 17.

Weeds of Queensland. Bindweed (Convolvulus arvensis ). Queensl. Agric. J. 33 :
322-323.

Note on Darlingia spectatissima F.v.Muell. with description of a new variety. J. Arnold
Arb. 11 : 231.

Queensland Vegetation in Handbook for Queensland. Austral. Ass. Adv. Sci., Brisbane
Meeting, 1930.

A new genus of Proteaceae from North Queensland. Kew Bull. 1930 : 234-235.

1931.

Two previously undescribed Queensland Myrtaceae. Proc. Roy. Soc. Queensl. 43 :
15-16.

Herbert Bennett Williamson. J. Bot. 69 : 172.

1932.

Two previously undescribed Rutaceae from south-eastern Queensland. Proc. Roy.
Soc. Queensl. 43 : 46-48.

Gustav Weindorfer. J. Bot. 70 : 261.

A previously undescribed Papuan Dipterocarp. Proc. Roy. Soc. Queensl. 43 : 49.

CYRIL TENISON WHITE.

Botanising in Tasmania. Queensl. Nat. 8 : 38-40.

The eucalypts or gum trees of the Brisbane district. [IX.] Queensl. Nat. 8 : 52-54.

The occurrence of the genus Aceratium (Elaeocarpaceae) in Australia. Kew Bull. 1932 :
42.

1933.

The genus Pleiogynium in Papua. Proc. Roy. Soc. Queensl. 45 : 27-28.

The eucalypts or gum trees of the Brisbane district. [X.] Queensl. Nat. 8 : 80-81.

Ligneous plants collected for the Arnold Arboretum in north Queensland by S. F.
Kajewski in 1929. Contrib. Arnold Arb. 4. 113 pp., 9 pi.

A plant poisonous to live stock. Queensl. Agric. J. n.s. 40 : 143-144. (With J. A.
Rudd.)

The noogoora burr ( Xantliium pungens ). A weed poisonous to livestock. Queensl.
Agric. J. n.s. 40 : 413.

Nomenclature of some Australian and Philippine Island plants. Kew Bull. 1933 : 45-4G.

1934.

Botany, Mount Edwards and district. (A) General. Queensl. Nat. 9 : 36-37.

The eucalypts or gum trees of the Brisbane district. [XI.] Queensl. Nat. 9 : 8-9.

The eucalypts or gum trees of the Brisbane district. [XII.] Queensl. Nat. 9 : 31-32.

Dendrobium (§ Rhizobium) Grimesii White & Summerhayes. Kew Bull. 1934 : 106,
(With V. S. Summerhayes.)

Queensland grasses. Queensl. Agric. J. n.s. 41 : 54-58.

Vinca rosea. A reputed cure for diabetes. Queensl. Agric. J. n.s. 41 : 598-600,

Queensland weeds. Blue weed or paterson’s curse ( Ectiium plantagineum). Queensl.
Agric. J. n.s. 41 : 304-305.

Queensland weeds. Mist flower ( Eupatorium riparium), Queensl. Agric. J. n.s. 41 :
519-521.

Queensland weeds. Gomphrena weed (Gompfirena decumbens). Queensl. Agric. J. n.s.
42: 83-84.

Queensland weeds. Nut grass (Cy perns rotundus). Sour grass or yellow grass (Paspalum
conjugatum). Queensl. Agric. J. n.s. 42 : 360-363.

Queensland weeds. Khaki weed ( Alternanfhera repens), Queensl. Agric. J. n.s. 42 :
583-584.

Thomas Lane Bancroft. J. Bot. 72 : 141-142.

1935.

Notes on the genus Ptychosperma in Queensland. Proc. Roy. Soc. Queensl. 46 : 80-82.
The structure and classification of Queensland orchids. Queensl. Nat. 9 : 55-61.

The common bracken ( Pteridium aquilinum). Queensl. Agric. J. n.s. 43 : 150-151.
Two trees of the genus Dysoxylum in North Queensland. North Queensl. Nat. 3: 34-36.

Lantana ( Lantana camara) and poison peach ( Trema aspera). Their effects on stock.
Queensl. Agric. J. n.s. 43 : 369-373. (With K. S. McIntosh.)

Queensland weeds. Indian jujube or china apple (Zizyphus mauritiana). Queensl,
Agric. J. n.s. 44 : 98-99.

Weeds of Queensland. Creeping knapweed ( Centaurea repens). Queensl. Agric. -J. n.s,
44 : 702-703.

1936.

Contribution to the Queensland flora, No. 5. Proc. Roy. Soc. Queensl. 47 : 51-84.

The eucalypts or gum trees of the Brisbane district. [XIII.] Queensl. Nat. 9 : 114-115.

Queensland weeds. Tridax daisy or tridax. ( Tridax procumbens, family Compositae)
Queensl. Agric. J. n.s. 45 : 258-259.

Groundsel bush or tree groundsel ( Baeeharis halimifolia). Queensl. Agric. J. n.s. 45 :
575-576.

Clovers and trefoils in Queensland. Queensl. Agric. J. n.s. 46 : 51-54.

Principles of botany for Queensland Farmers. Queensl. Agric. J. n.s, 4.6 : 196-228,

358-391, 481-505, 602-632, 747-766.

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

1937.

Principles of botany for Queensland farmers. Queensl. Agric. J. n.s. 47 : 17-48, 154-180.

Queensland weeds. Johnson grass and wild sorghum. Queensl. Agric. J. n.s. 47 :
365-368.

Is button grass poisonous to sheep ? Queensl. Agric. J. n.s. 47 : 502.

Pitted blue grass. A pest in coastal pastures. Queensl. Agric. J. n.s. 48 : 70.
Hexham scent. Queensl. Agric. J. n.s. 48 : 430.

Two new dendrobs for north Queensland. Queensl. Nat. 10 : 25-26. (With IT. M. It.
Rupp.)

The Australian species of RJiodamnia. Blumea Suppl. 1 : 214-218.

1938.

The eucalypts or gum trees of the Brisbane district. [XIV.] Queensl. Nat. 10 : 72-73.
Queensland weeds. Anoda cristata. Queensl. Agric. J. n.s. 49 : 332-333.

Queensland weeds. Shrubby or upright mist flower (Eupatorium adenophorum). Queensl.
Agric. J. n.s. 50 : 8-9.

The varieties of guinea grass cultivated in Queensland. Queensl. Agric. J. n.s. 49 :
110-112.

Bindweed — A serious weed pest. Queensl. Agric. J. n.s. 50 : 7.

Shrubby or upright mist flower. Queensl. Agric. J. n.s. 50 : 83.

Two weeds poisonous to stock. Queensl. Agric. J. n.s. 50 : 217.

A cause of shivers or staggers in stock. Queensl. Agric. J. n.s. 50 : 533.

Melilot or Hexham scent. Queensl. Agric. J. n.s. 50 : 661-662.

Principles of botany for Queensland farmers. Govt. Printer, Brisbane. 232 pp., 102
pi. (Previously published in parts in 1936-7 in Queensl. Agric. J.)

Ferns of Mt. Spurgeon, North Queensland. Viet. Nat. 54: 147-151. (With D. A. Goy).
John Fredei'ick Bailey. J. Bot. 76 : 307.

1939.

Contributions to the Queensland Flora, No. 6. Proc. Roy. Soc. Queensl. 50 : 66-87.
Solarium hispidum Pers. : its distribution and synonymy. Kew Bull. 1939 : 666-668.
Wild sunflower — A poisonous plant. Queensl. Agric. J. n.s. 51 : 97-98.

Hoary cress ( Lepidium draba). A possible serious weed pest in Queensland. Queensl.
Agric. J. n.s. 52 : 658-661.

1940.

A new type of sundew from north Queensland. Viet. Nat. 57 : 94-95.

A new Dendrobium from New Guinea. Austr. Orchid Rev. 5 : 74-75.

A history of Australian orchids (in part). Austr. Orchid Rev. 5 : 89.

1941.

A new genus of Flacourtiaceae (Pangieae-Hydnocarpinae) from tropical Queensland.
J. Arnold Arb. 22: 143-144.

Red-flowered lotus or bird’s foot trefoil ( Lotus coccineus). Queensl. Agric. J. n.s. 55 :
297-299. (With E. H. Gurney.)

Shade and ornamental trees and shrubs for the pig farm. Queensl. Agric. J . n.s. 55 :
300-302. (With E. J. Sheldon.)

Noxious weeds. Queensl. Agric. J. n.s. 56 : 332-336.

Myoporum acuminatum (strychnine bush) — A plant poisonous to stock. Queensl.
Agric. J. n.s. 56 : 124-125.

Myoporum acuminatum : A plant poisonous to stock. Austr. Vet. J. 17 : 104-105.
(With J. Legg.)

A new variety of Dendrobium from N. Queensland. Austr. Orchid Rev. 6 : 53.
Queensland Pasture Plants. Pamphlet No. 85. Queensl. Dept. Agric. & Stock. 29 pp.
(With S. L. Everist and C. W. Winders.)

CYRlL TRXJSOX \V II [TE.

4 i

1942.

Contributions to the Queensland Flora, No. 7. Proc. Roy. Soc. Queensl. 53 : 201-228.
A rare lycopod from the Lamington National Park, S.E. Queensland. Queensl. Nat.
12: 32-33.

Some Papuan Myrtaeeae. J. Arnold Arb. 23 : 79-92.

The genus Dendrobium in Australia. Austr. Orchid Rev. 7 : 4-7.

1943.

On two interesting species of Marsdenia (family Asclepiadaceae). Queensl. Nat. 12 :
54-55.

Has Dendrobium opfiioglossum been rediscovered ? Austr. Orchid Rev. 8 : 19.

The genus Dendrobium in New Guinea and the Solomon Islands. Austr. Orchid Rev.
8 : 62-63.

1944.

Contributions to the Queensland Flora, No. 8. Proc. Roy. Soc. Queensl. 55 : 59-83.
The algaroba bean or mesquite as a pest plant. Queensl. Agric. J. n.s. 58 : 360-361.
Gomphrena weed. Queensl. Agric. J. n.s. 58 : 179-180.

The sword bean ( Canavalia gladiata). Queensl. Agric. J. n.s. 58 : 307.

Crofton weed, a serious pest. Queensl. Agric. J. n.s. 59 : 154-155.

New Guinea timbers. Queensl. Agile. J. n.s. 59 : 278-282.

Giant sensitive plant. A very serious weed pest in North Queensland. Queensl. Agric.
J. n.s. 59 : 341-342.

Cretan weed. Queensl. Agric. J. n.s. 59 : 343.

A new variety of Dendrobium plialaenopsis (Fitzg.). Austr. Orchid Rev. 9 : 35.

1945.

Try on memorial lecture : Henry Try on — First Hon. Secretary, Royal Society of

Queensland, and his place in Queensland Science. Proc. Roy. Soc. Queensl.
56 : 77-80.

Wattles or Acacias of the Brisbane district. Queensl. Nat. 12 : 107-113.

Chinese spinach. A useful summer vegetable. Queensl. Agric. J. n.s. 60 : 83-84.
Bulbous oat grass or onion couch, a possible weed pest. Queensl. Agric. J. n.s. 60 : 154.

Shade and shelter plants for the pig farm. Queensl. Agric. J. n.s. 60 : 226-227.

(With E. J. Shelton.)

Chicory — A weed pest often confused with skeleton weed. Queensl. Agric. J. n.s. 60 :
352.

The Bailey family and its place in the botanical history of Australia. J. Hist. Soc.
Queensl. 3 : 362-368.

Flora of Melanesia. Queensl. Geogr. J. n.s. 49 : 64-67.

1946.

Contributions to the Queensland flora, No. 9. Proc. Roy. Soc. Queensl. 57 : 21-36.

An important contribution to our knowledge of Dutch New Guinea (Fragmenta Papuana
(Observations of a naturalist in Netherlands New Guinea) by H. J. Lam ; a
review and commentary). Queensl. Nat. 13 : 13-18.

St. Barnaby’s Thistle. Queensl. Agric. J. n.s. 62 : 271-272.

Dr. Eugen Hirschfeld : An appreciation. Queensl. Agric. J. n.s. 63 : 117-118.

The devil’s fig ( Solanum torvum). Queensl. Agric. J. n.s. 63 : 280-281.

The Australian species of Antirhea , and a new name for a Cuban species. J. Arnold
Arb. 27 : 121-122.

Papuodendron, a new genus of arborescent Malvaceae from New Guinea. J. Arnold
Arb. 27 : 272-274.

A new species of Longetia : the botanical identity of a “ pink cherry ” of Dorrigo
timber-getters. Proc. Linn. Soc. N. S. Wales 71 : 236-238. (With W. A. W.
de Beuzeville.)

Botanizing in the British Solomon Islands. Austr. J. Sci. 9 : 62-64.

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

1947.

Notes on two species of Araucaria in New Guinea and a proposed new section of the
genus. J. Arnold Arb. 28 : 259-260.

1948.

Three species of Endiandra (family Lauraceae) from eastern Australia. Proo. Boy.
Soc. Queensl. 59 : 151-152.

A new species of Austrobaileya (Austrobaileyaceae) from Australia. J. Arnold Arb.
29 : 255-256.

1949.

The genus Embothrium Forst. (family Proteaceae) in Australia. Proe. Roy. Soc.
Queensl. 60 : 43-44.

Einschia — A genus of “ nut ” trees of the south-west Pacific. Pacific Sci. 3 : 187-194.
Star thistle — a new weed pest. Queensl. Agric. J. n.s. 69 : 84-85.

Wild cottons — declared noxious weeds. Queensl Agric. J. n.s. 69 ; 143-145.

Weir vine — a declared noxious plant. Queensl. Agric. -J. n.s. 69 : 143-145.

1950.

Additions to the flora of Arnhem Land. Proc. Roy. Soc. Queensl. 61 : 55-58.

F. M. Bailey : His life and work. Proc. Roy. Soc. Queensl. 61 : 105-114.

Ligneous plants from the Solomon Islands (and New Guinea). J. Arnold Arb. 31 :
81-116.

Report on the botany of the Mt. Alford excursion. Queensl. Nat. 14 : 35-36.

1951.

Some noteworthy Myrtaceae from the Moluccas, New Guinea and the Solomon Islands.
J. Arnold Arb. 32 : 139-149.

A previously undescribed Pomaderris from south-east Queensland. Queensl. Nat. 14 :
48"

VOL. LXII., No. 4.

REDUCTIONS IN ELAEOCARPUS.

By E. D. Merrill, Arnold Arboretum, Harvard University.

(Issued separately, 15th August, 1952.)

SUMMARY.

90 published names under Elaeocarpus are reduced to synonymy.

Elaeocarpus Linn, is a genus characteristic of the Old World tropics, a
few species only occurring in subtropical and even in subtemperate regions.
The generic range is tropical Africa and Madagascar to Japan, southward
through Malaysia to eastern Australia, New Caledonia and New Zealand,
eastward to Hawaii. Speciation within the limits of the Malay
Archipelago (including the Philippines and Papuasia) is remarkable. As in
all large genera in this vast region that are characteristic of the primary
forests, there are a few widely distributed and more or less variable species,
but everywhere, particularly in the insular areas, there is a very highly
developed local endemism ; and a high percentage of these local endemics
are very sharply defined species.

Up to 1900 about 220 binomials had been published under Elaeocarpus ,
yet Bentham and Hooker f., in 1862, most conservatively estimated the
number of distinct species as about 50 ; and 28 years later K. Schumann
surmised that the number might be more than 60. In the past 50 years
about 330 new Elaeocarpus binomials have been published, mostly for
optimistically proposed 4 4 new species.” Anyone now attempting to work in
this particular group is faced with approximately 550 published binomials
under Elaeocarpus alone.

It was clear that in the proposals made in the past 50 years there was a
considerable amount of duplication of work, due to one cause or another, but
chiefly, it is suspected, because a systematist working in one place, be it
Buitenzorg, Singapore, Manila, London, Berlin, or any other taxonomic
center, did not have access to authentically named specimens representing
many of the proposed species. The accepted priority and homonym rules
are also involved. There is evidence that sometimes ample validly published
descriptions were not critically studied, and in at least one case, that it
was not even scanned. The situation is further complicated because too
many of the proposed and described “ new species ” were based on fruiting
specimens, or on those where only juvenile flower buds were present. Added
to the confusion is the unsatisfactory work of Knuth*, who between the
years 1938 and 1941, described no less than 80 new species without in any
case indicating the subgenus or section, and rarely suggesting specific
alliances ; twenty- two of these eighty species are herein reduced to synonymy
on the basis of an actual study of excellent isotype material in each case.
The Knuth holotypes were all destroyed in the Berlin disaster in 1943.

Fortunately, during the summer of 1949, I was able to examine most
of the actual types of species described by Blume and by Miquel in the
Leiden and Utrecht herbaria, and those of the British botanists preserved
at Kew. I have, hence, in practically every case, been able to reach my
conclusions on the basis of an actual examination of the types, isotypes,

* Knuth, R. Elaeocarpaceae novae. Decades 1-2, Repert. Sp. Nov. 44 : 124-132
(1938); Decades 3-4, op. cit. 48 : 72-79 (1940); Decades 5-6, op. cit. 49 : 66-73 (1940);
Decades 7-8, op. cit. 50 : 81-88 (1941).

50 PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

and other historical specimens. All of the original, and many of the repeated
descriptions have been studied. In difficult groups which need a thorough
and critical revision on the basis of very ample collections, such as the
Elaeocarpus sphaericus (Gaertn.) K. Schum. and E. stipularis Blume
complexes, some future monographers may not agree with all of my proposed
reductions. I am, however, confident that in the vast majority of the
cases listed below, the indicated reductions will be upheld.

The net result of my studies to date is the reduction of 90 proposed
binomials to synonymy; 17 of these were published before 1900, the others
within the past fifty years. It is evident, as studies and comparisons are
continued, that still other reductions will inevitably be made. Yet at the
same time one will find in all of the large herbaria a certain number of
sharply defined species which have never been studied, and which remain
unnamed and undescribed. The number of these increases as collections
come in from previously inadequately explored regions, and especially from
those large areas which have not previously been visited by a botanist or
even a botanical collector.

E. acuminatus Koord. & Val. Meded. Lands Plant. 11 : 258 (1894) (Bijdr*
1 : 258), non Wall. (1830, 1874) = E. oxypyren Koord. & Val. op-
cit. 33 : 419 (1900) ; Ic. Bogor. 3 : pi 420 (1914). Java.

E. aemulus A. C. Sm. in Jour. Arnold Arb. 25 : 232 (1944) = E. multi-
scissus Knuth in Repert. Sp. Nov. 50 : 86 (1941). Clemens 5019
typifies both. New Guinea.

E. argyroides Hance in Jour. Bot. 15 : 330 (1877) = E. griffithU
(Wight) A. Gray ; see Merrill in Jour. Arnold Arb. 19 : 50 (1938).
Indo-China ; range Siam, Malay Peninsula, Sumatra, Borneo.

E. Augustus Knuth in Repert. Sp. Nov. 44 : 125 (1938) = E.

dolichobotrys Merr. in Jour. Straits Br. Roy. As. Soc. 77 : 198
(1917). Borneo.

E. baclayanensis Elm. Leafl. Philip. Bot. 4 : 1188 (1911) |§ E.

sphaericus (Gaertn.) K. Schum. fide Van Steenis in Bull. Jard. Bot.
Buitenz. Ill, 17 : 409 (1948). Philippines ; range India to New
Guinea and Samoa, i.e., sensu lat.

E. baramensis Knuth in Repert. Sp. Nov. 44 : 126 (1938) = E. ferevipes
Merr. in Jour. Straits Br. Roy. As. Soc. 86 : 327 (1922) ; see Airy
Shaw in Kew Bull. 1949 : 165 (1949). Borneo.

E. barbulatus Knuth, l.c. 44 : 126 (1938) == E. nitidus Jack (1920) ; see
Airy Shaw, 1. c. Borneo ; range Malay Peninsula, Sumatra.

E. bataanensis Merr. in Philip. Jour. Sci. 10 : Bot. 41 (1915) = E.
calomala (Blanco) Merr. Philippines.

E. bellus Knuth in Repert. Sp. Nov. 49 : 67 (1940) j| E. merrittii Merr. in
Philip. Jour. Sci. 2 : Bot. 280 (1907). Merrill 5582 typifies both.
Philippines.

E. borneensis Knuth, op. cit. 44 : 127 (1938) =-E. ferrugineus (Jack)
Steud. ; see Ridley in Kew Bull. 1938 : 234 (1938), and Airy Shaw,
op. cit. 1949 : 165 (1949). Borneo ; range Malay Peninsula,

Sumatra.

REDUCTIONS IN ELAEOCARPUS.

E. beccarii Warb. in Repert. Sp. Nov. 18 : 328 (1922), non Aug. DC.
(1903) == E. euneuras Stapf ex Ridl. in Kew Bull. 1938 : 234 (1938).
Borneo.

E. brachyphyllus Kiiuth, op. cit. 50 i 82 (1941) = E. hainanensis
Oliv. in Hook. Ic. 25 : pi. 2462 (1896). ( E . hainanensis Oliv. var.
brachyphyllus Merr. in Lingnan Sci. Jour. 5 : 123 (1927). Hainan ;
range Kwangtung, Indo-China, Siam, Malay Peninsula.

E. celebesianus Baker f. in Jour. Bot. 62 : Suppl. 14 (1924) = E.
teijsmannii Koord. & Val. in Koord. Meded. Lands Plant. 19 : 368
(1898), nom., Ic. Bogor. 2 : pi. 128 (1904), descr. ; see Van Steenis
in Bull. Jard. Bot. Buitenz. III. 18 : 210 (1949). Celebes.

E. clemensiae Knuth in Repert. Sp. Nov. 44 : 128 (1938) — E. dementis
Merr. in Jour. Straits Br. Roy. As. Soc. 77 : 195 (1917). The basic
specimens are Clemens 27 66, 51312. Borneo.

E. dallasensis Knuth, op. cit. 49 : 69 (1940). — E. elliptifolius Merr.
op. cit. 193 (1917). The basic specimens are Clemens 10783, 26231.
Borneo.

E. decorus A. C. Sm. in Jour. Arnold Arb. 25 : 242 (1944) = E.
densifloras Knuth, op. cit. 50 : 82 (1941). Clemens 2420 typifies
both. New Guinea.

E. decurvattjs Diels in Notizbl. Bot. Gart. Berlin 11 : 214 (1931) = E«
varunna Buch.-Ham. in Wall. List. no. 2666 (1830), nom., Mast, in
itook. f. FI. Brit. Ind. 1 : 407 (1874), descr. China (Kwangsi) ; range
Southern China and Indo-China to Burma and Northern India.

E. dioicus Turcz. in Bull. Soc. Nat. Mosc. 19 (2) : 493 (1846) = E.
japonicus Sieb. and Zucc. (1845). Japan ; range central, western
and southern China.

E. ellipticus Nakai in Bot. Mag. Tokyo 18 : 67 (1904), non Sm. (1809) =
E. decipiens Hemsl. in Jour. Linn. Soc. Bot. 23 : 95 (1886) ; see
Metcalf in Sunyatsenia 6 : 178 (1941). Japan ; range Riu Kiu
Islands, Formosa.

E. fagaceus Knuth in Repert. Sp. Nov. 44 : 128 (1938) = E. cupreus
Merr. in Jour. Straits Br. Roy. As. Soc. 77 : 193 (1917). Borneo.

E. ferrugineus Bedd. FI. Sylv. South. Ind. pi. 112 (1869), non E.
ferrugineus (Jack) Steud. (1840) = E. recurvatus Corner in Gard.
Bull. Straits Settl. 10 : 319, 325 (1939). India.

E. FissiSTiPULUS Miq. FI. Ind. Bat. 1 (2) : 210 (1859) =. E. stipuiaris
Blume Bijdr. 120 (1825), sensu lat. Sumatra ; range Malay Peninsula,
Sumatra, Java, and Borneo.

E. fusicarpus Elm. Leaf!. Philip. Bot. 4: 1174 (1911) =E. procerus
Aug. DC. in Elm. Leaf!. Philip. Bot. 2 : 636 (1909) ; see Airy Shaw
in Kew Bull. 1949 : 165 (1949). Philippines.

E. gambir Becc. Nelle Foreste di Borneo 594 (1902), nom. = E. stipuiaris
Blume (1825), sensu lat. Borneo ; range Malay Peninsula, Sumatra,
Java.

52 PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

E. glabripetalus Merr. in Philip. Jour. Sci. 21 : 501 (1922) = E.
sylvestris (Lour.) Pers. Kwangtung ; range southern China, Hainan,
and Indo-China.

E. gracilipes Knuth in Repert. Sp. Nov. 49 : 70 (1940) = E. nitidus
Jack Mai. Misc. 1 (5) : 41 (1820). Borneo ; range, Malay Peninsula,
Sumatra.

E. henryi Hance in Jour. Bot. 28 : 322 (1885) = E. sylvestris (Lour.)
Pers. China, Kwangtung ; range, southern China and Indo-China.

E. holosericeus Blume ex Koord. & Val. in Meded. Lands Plant. 33 :
422 (1900) (Bijdr. 5 : 422) — E. obtusus Blume Bijdr. 123 (1825) ; see
Adelbert in Blumea 6 : 312(1949). Java; range Sumatra, Borneo.

E. hosei Merr. in Jour. Straits Br. Roy. As. Soc. 11 : 197 (1917) = E.
glaber Blume Bijdr. 122 (1825). Borneo ; range Sumatra, Java.

E. hullettii King in Jour. As. Soc. Bengal 60 (2) : 132 (1891) (Mater.
FI. Malay Penin. 1 : 242) = E. palembanicus (Miq.) Corner in Gard.
Bull. Straits Settl. 10 : 323 (1939). Malay Peninsula ; range

Sumatra.

E. integerrimus Lour. FI. Cochinch. 338 (1790) ,= Qchna mtegerrima
(Lour.) Merr. in Trans. Am. Philos. Soc. II. 24 (2) : 265 (1935)
( Discladium harmandii Van Tiegh. ; Ochna harmandii H. Lecomte).
Indo-China ; range Hainan.

E. isotrxchus F.-Vill. Novis. App. FI. Filip. 31 (1880) (Monocera isotricha
Turcz.) = E. calomala (Blanco) Merr. in Philip. Jour. Sci. 10 : Bot. 43
(1915). Philippines.

E. kajewskii Guillaumin in Jour. Arnold Arb. 12 : 232 (1931)

= Aceratinm braithwaitei (F. Muell.) Schltr. in Bot. Jahrb. 54 : 103
(1916) ( Aristoteleia braithwaitei F. Muell. in South. Sci. Record 1 : 149
(1881). The type of F. Mueller’s species was from Tanna, that of
Guillaumin’s from Aneityum, New Hebrides, and both descriptions
apply to a single species.

E. kwangtungensis Hu in Jour. Arnold Arb. 5 : 2^9 (1924), Contr.
Biol. Lab. Sci. China 1 (2) : 3 (1925) = E. sylvestris (Lour.) Pers.
Kwangtung ; range southern China and Indo-China.

E. lacex Craib in Kew Bull. 1925 : 23 (1925) = E. liainanensis Oliv.
(1896) fide Gagnepain in Lecomte FI. Gen. Indo-Chine Suppl. 1 : 478
(1945). Siam ; range Hainan, Indo-China, Malay Peninsula.

E. lagtjnensis Knuth in Repert. Sp. Nov. 50 : 85 (1941) = E. sessilis
Knuth, op. cit. 87. Philippines.

E. leftopus A. C. Sm. in Jour. Arnold Arb. 25 : 234 (1944) = E.
albiflorus Knuth in Repert. Sp. Nov. 50 : 81 (1941). Clemens 1022
typifies both. New Guinea.

E. leptostachys Wall. List no. 2672 (1830), nom., C. Muell. Anot. Fam.
Elaeoc. 23 (1849), descr. = E. tectorius (Lour.) Merr. North-eastern
India ; range Burma to Indo-China and the Malay Peninsula.

E. leptomischus Ridl. in Jour. Fed. Malay States Mus. 5 : 30 (1913)
= E. palembanicus (Miq.) Corner in Gard. Bull. Straits Settl. 10 : 323
(1939). Malay Peninsula ; range Sumatra.

REDUCTIONS IN ELAEOCARPUS.

E. leytensis Merr. in Philip. Jour. Sci. 8: Bot. 383 (1913) = E. multifloras
(Turcz.) F.-Vill. Novis. App. FI. Filip. 31 (1880) ( Monocera multiflora
Turcz. in Bull. Soc. Nat. Mosc. 19 (2) : 495 (1846). Philippines.

E. linearifolius Knuth in Repert. Sp. Nov. 49 : 66 (1940) = E.

hainanensis Oliv. (1896). Indo-China ; range Kwangtung, Hainan,
Siam, Malay Peninsula.

E. longifolius Blume var. fusiformis Corner in Gard. Bull. Straits
Settl. 10 : 321 (1939) = E. submonoicus Miq. FI. Ind. Bat. Suppl.
407 (1862). Malay Peninsula, Borneo ; range Sumatra.

E. longipetiolatus C. T. White in Bot. Bull. Queensl. Dept. Agr. 21 : 3
(1919), non Merr. (1917) = E. micliaelii C. T. White in Kew Bull.
1933 : 45 (1933). Queensland.

E. luridus Stapf ex Ridl. in Kew Bull. 1938 : 232 (1938) = E.

pacliyoplirys Warb. in Repert. Sp. Nov. 18 : 327 (1922). Borneo.

E. maclurei Merr. in Lingnan Sci. Jour. 13 : 63 (1934) = E. limitaneus
Hand.-Maz. in Sinensia 3 : 193 (1933). Hainan ; range Southern
China, Indo-China.

E. macrocarpus Ridl. FI. Malay Penin. 1 : 32 (1922) = E. robustus
Roxb. (1814, 1832) fide Corner in Gard. Bull. Straits Settl. 10 : 325
(1939) = E. tectorius (Lour.) Merr. in Trans. Am. Philos. Soc. II.
24 (2) : 256 (1935) (Craspedum tectorium Lour. FI. Cochinch. 336
(1790).) Malay Peninsula ; range north-eastern India to Indo-China
and the Malay Peninsula.

E. macropus Warb. ex Knuth in Repert. Sp. Nov. 48 : 76 (1940) = E.
petiolatus (Jack) Wall. Celebes ; range Indo-China and Siam to
the Malay Peninsula, Sumatra and Borneo.

E. magnifolius Knuth in Repert. Sp. Nov. 44 : 129 (1938), non

Christoph. (1935) = E. gustaviifolius Knuth, op. cit. 50 : 84 (1941),
Clemens 29481, in flower, 28078, in fruit. Borneo.

E. m akin oi Kaneh. & Hatus. in Rept. Forest. Exp. Sta. Kyushu Univ.
3 : 119 (1933), Formosan Trees, ed. 2, 433 (1936), in syn. —E. ellipticus
Nakai = E. japonicus Sieb. & Zucc. Formosa ; range central and
southern China, Indo-China.

E. maquilingensis Elm. Leafl. Philip. Bot. 8 : 3080 (1919) = E. pendulus
Merr. in Govt. Lab. Publ. 29 : 27 (1905). Philippines.

E. matangensis Knuth in Repert. Sp. Nov. 44 : 130 (1938) = E.
pedunculatus Wall. List no. 2678 (1830), nom. ; Mast, in Hook. f.
FI. Brit. Ind. 1 : 408 (1874), descr. Borneo ; range Malay Peninsula,
Sumatra.

E. megacarpus Elm. Leafl. Philip. Bot. 7 : 2627 (1915) = E. monocera
Cav. Ic. 6 : pi. 501 (1801). Philippines.

E. megacarpus Schltr. in Bot. Jahrb. 54 : 131 (1916), non Elm. (1915)
= E. schlechteriamis A. C. Sm. in Jour. Arnold Arb. 25 : 256 (1944).
New Guinea.

E. merrillii Elm. ex Merr. Enum. Philip. FI. PL 3 : 14 (1923), nom. in
syn., Leafl. Philip. Bot. 10 : 3729 (1939), descr. anglice = E.

argenteus Merr. in Govt. Lab. Publ. 23 : 26 (1905). Philippines.

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

E. micranthus Vieill. in Bull. Soc. Linn. Normandie 9 : 392 (1865) == E .
rotundifolius Brongn. & Gris in Bull. Soc. Bot. France 8 : 261
(1861), fide Guifiaumin in Not. Syst. 2 : 92 (1911). New Caledonia.

E. microphyllus Warb. in Repert. Sp. Nov. 18 : 328 (1922), non Elm.
(1911) = E. octantherus Aug. DC. in Bull. Herb. Boiss. IX. 3 : 367
(1903). Borneo.

E. miquelii Hochr. PL Bogor. Exsic. 27 (1904), excl. spec. cit. = E,
obtusus Blume Bijdr. 123 (1925). Java ; range Malay Peninsula,
Sumatra, Borneo. (No. 50Hochreutiner, PI. Bogor. Exsic., is actually

E. pedunculatus Wall., not at all the species that Miquel described
and which Hochreutiner renamed.)

E. molloides Knuth in Repert. Sp. Nov. 49 : 71 (1940) == E. canipes
Knuth, op. cit. 44 : 127 (1938). Clemens 20039, 32147, one in flower,
one in fruit. Borneo.

E. MYRTILLUS Schltr. in Bot. Jahrb. 39 : 180 (1906) = E. vaccinioides

F. Muell. ex Brongn. in Bull. Soc. Bot. France 12 : 301 (1865). See
Guillaumin in Not. Syst. 2 : 94 (1911). New Caledonia.

E. nigropunctatus Merr. in Jour. Straits Branch Roy. As. Soc. 77 : 196
(1917) = E. nitidus Jack (1820) ; see Airy Shaw in Kew Bull. 1949 :
165 (1949). Borneo ; range Sumatra, Malay Peninsula.

E. nutans Knuth, op. cit. 44 : 131 (1938) — E. tomentosus Blume

Bijdr. 121 (1825) = E. stipularis Blume, 1. c., sensu lat. Borneo ;
range Malay Peninsula, Sumatra, Java.

E. ochraceus Ridl. in Trans. Linn. Soc. II. Bot. 9 : 21 (1916) =
Aceratium ocliraceum Schltr. in Bot. Jahrb. 55 : 194 (1918). New
Guinea.

E. ochraceus Stapf ex Ridl. in Kew Bull. 1938 : 230 (1938), non Ridl.
(1916) = E. eupreus Merr. in Jour. Straits Branch Roy. As. Soc.
77 : 193 (1917). Borneo.

E. omeiensis Rehd. & Wils. in Sargent PI. Wils. 2 : 260 (1915) = E.
sylvestris (Lour.) Pers. Szechuan ; range central and southern China
and Indo- China.

E. ovalifolius Wall. List no. 2665b (1830), nom., C. Muell. Anot. Fam.
Elaeoc. 21 (1849), descr. = E. tectorius (Lour.) Merr. India ; range
Burma to Indo-China and the Malay Peninsula.

E. oxyadenius Warb. in Repert. Sp. Nov. 18 : 327 (1922) == E. beccarii
Aug. DC. in Bull. Herb. Boiss. II. 3 : 367 (1903). Beccarii 540
typifies both. Borneo.

E. papuanus Knuth in Repert. Sp. Nov. 50 : 86 (1941) = E. brassii
Knuth, op. cit. 48 : 73 (1940) ; see A. C. Smith in Jour. Arnold
Arb. 25 : 240 (1944). New Guinea ( Brass 7450, 7566).

E. patens Knuth, op. cit. 48 : 77 (1940) = E. viseosus Warb. in Bot.
Jahrb. 18 : 201 (1893) ; see A. C. Sm. in Jour. Arnold Arb. 25 : 246
(1944). New Guinea.

E. philippinensis Warb. in Perk. Frag. FI. Philip. 100 (1904) = E.
caiomala (Blanco) Merr. ; see Merr. Enum. 3 : 15 (1923) for

synonymy. Philippines.

REDUCTIONS IN ELAEOCARPUS. 55

E. polycarpus Stapf ex Ridl. in Kew Bull. 1938 : 230 (1938) (Aug. 15)
= E. multinervosus Knuth in Repert. Sp. Nov. 44 : 130 (1938)
(June 30). Beccari 2698 typifies both. Borneo.

E. polychistus Schltr. in Bot. Jahrb. 40 : Beibh 92 : 28 (1908; ==
E. persiciiolins Brongn. & Gris in Bull. Soc. Bot. France 8 :
202 (1861). New Caledonia.

E. quercifolius Gagnep. in Not. Syst. 1 : 124 (1910), non Baker (1883)
= E. viguieri Gagnep., op. cit. 3 : 133 (1915). Xndo-China.

E. ramosii Knuth in Repert. Sp. Nov. 44 : 131 (1938) = E. floribundus
Blume Bijdr. 120 (1925). Borneo ( Ramos 1227) ; range British
India to Indo-China southward to Sumatra, Java, and Borneo.

E. rejangensis Knuth, l.c. J= E. brevipes Merr. (1922) ; see Airy Shaw
in Kew Bull. 1949 : 165 (1949). Borneo.

E. reticulatus Ridl. in Jour. Straits Br. Roy. As. Soc. 61 : 2 (1912), non
Sm. (1809) — E. nanus Corner in Gard. Bull. Straits Settl. 10 : 322
(1939). Malay Peninsula.

E. rivularis Vieill. ex Britten in Forbes, Nat. Wand. East. Archipel. 501
(1885), nom. ; Guillaumin in Not. Syst. 2 : 92 (1911), nom. ; Baker
f. in Jour. Bot. 62 : Suppl. 14 (1924), nom. in nota, non Gagnep.
(1910) = E. rotmidifolius Brongn. & Gris in Bull. Soc. Bot. France
8 : 201 (1861). New Caledonia.

E. robustus Roxb. Hort. Beng. 42 (1814), nom., G. Don Gen. Syst.
1 : 559 (1831), descr., Roxb. FI. Ind. ed. 2, 2 : 597 (1832), descr „
= E. tectorius (Lour.) Merr. in Trans. Am. Philos. Soc. II. 24 (2) :
256 (1935) ( Dicer a craspedum J. F. Gmel. ex DC. Prodr. 1 : 520'
(1824) ; Craspedum tectorium Lour. FI. Cochinch. 336 (1790)). India ;
range Burma, Siam, Indo-China, the Malay Peninsula and Sumatra.
See, however, Corner in Gard. Bull. Straits Settl. 10 : 325 (1939)
for a rather strange dissenting opinion, considering that Loureiro’s
type is still extant and has been examined.

E. rumphii Merr. Interpret. Herb. Amb. 347 (1917) = E. erenatus (Raf.)
Merr. Jour. Arnold Arb. 29 : 212 (1948) ( Ayparia crenata Raf.
Sylva Tellur. 154 (1838)). Amboina. An E. sphaericus (Gaertn.) K.
Schum., sensu lat. ?

E. s adikanen sis Knuth in Repert. Sp. Nov. 49 : 71 (1940) = E. petiolatus
(Jack) Wall. Borneo ; range Sumatra, Malay Peninsula, Indo-
China, Hainan.

E. scortechinii King in Jour. As. Soc. Bengal 60.(2) : 132 (1891) (Mater.
FI. Malay Penin. 1 : 233) = E. stipularis Blume, sensu lat. Malay
Peninsula ; range Sumatra, Borneo, Java.

E. sericeus Stapf in Trans. Linn. Soc. II. Bot. 4 : 137 (1894), non Baker
(1883) = E. congestiMius Knuth in Repert. Sp. Nov. 49 : 68 (1940).
The two species were independently published. Borneo (Mount-
Kinabalu).

E. stenophyllus Merr. in Pap. Michigan Acad. 19 : 168 (1934) - E.
salicifolius King in Jour. As Soc. Bengal 60 (2) : 125 (1891) (Mater.
FI. Malay Penin. 1 : 234). Sumatra ; range Malay Peninsula.

56 PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

E. subdenticulatus Miq. FI. Ind. Bat. Suppl. 408 (4862) = E. petiolatus
(Jack) Wall. Sumatra ; range Malay Peninsula, Indo-China,
Hainan, Borneo.

E. subglobosus Merr. in Philip. Jour. Sci. 4 : Bot. 290 (1909) ==• E.
sphaericus (Gaertn.) K. Schum., fide Corner in Gard. Bull. Straits
Settl. 10 : 326 (1939), et Van Steenis in Bull. Jard. Bot. Buitenzorg
III. 17 : 409 (1948). Philippines ; range ( sensu lat.) India to
Southern China through Malaysia to Samoa.

E. subsessilis Hand.-Maz. in Symb. Sin. 7 : 614, pi. 21, fig. 5 (1933) =
E. sylvestris (Lour.) Pers. Hunan, Fukien, Chekiang ; range western
and southern China and Indo-China.

E. valetonii Hochr. PI. Bogor. Exsicc. 29 (1904) = E. subpuherus
Miq. FI. Ind. Bat. Suppl. 406 (1862). Sumatra ; range Bangka,
Billiton, cultivated at Buitenzorg, Java (VI. C. 97) from which
apparently came no. 55 of the Hochreutiner exsiccatae.

E. versicolor Elm. Leaf!. Philip. Bot. 4: 1178 (1911) =E. cumingii
Turcz. in Bull. Soc. Nat. Mosc. 19 (2) : 491 (1846). Philippines.

E. yengtangensis Hu in Jour. Arnold Arb. 5 : 229 (1924) = E. japonicus
Sieb. & Zucc. (1845). China (Chekiang) ; range Japan to southern
and western China.

E. yentanensis Hu in Contr. Biol. Lab. Sci. Soc. China 1 (2) : 4 (1925) =
praec.

E. yunnanensis Brandis ex Tutcher in Rept. Bot. For. Dept. Hongkong
1914 : 229 (1915), nom., reprint, Chun in Sunyatsenia 1 : 170

(1933) = E. japonicus Sieb. & Zucc. (1845). Kwangtung, Yunnan ;
range Japan to western and southern China.

Vol. LXIL, No. 5.

VEGETATIVE HABIT IN THE GENUS
EULOPHIA (Orchidaceae).

By R. E. Holttum (University of Malaya, Singapore).

(. Issued separately, 15th August, 1952.)

SUMMARY.

The vegetative habit of the genus Eulophia is described, with particular reference
to E. bicarinata.

The genus Eulophia (including Lissochilus ) comprises some 300 species
distributed throughout the warmer parts of the Old World, mainly in
Africa. My acquaintance with it is limited to the six species occurring in
the Malay Peninsula, of which five are widely distributed, two with a range
from India to Australia. One of these species, which thus connects Malaya
and Queensland, has only recently been found for the first time in Malaya ;
it is E. bicarinata (also known as E. venosa). In vegetative form it is different
from the others, and appears to be of considerable interest. It has also a
connection with the late Mr. C. T. White, and the last letter he wrote me
was about a drawing of this Eulophia which he made many years ago.

In the various descriptions of the African species of Eulophia which
have been brought into cultivation, there is little definite information
about the way in which the rhizome and pseudobulbs develop. The best
source of information about vegetative structure is Pantling’s beautiful
plates of Sikkim orchids (King and Pantling, 1898). These drawings show
the great variety of vegetative habit in the Sikkim species. Some indication
of development is given, though this is not discussed in the text. These
plates include the best illustration of E. bicarinata that I have seen.

The origin of my interest in this matter was the receipt of an inflorescence
of a Eulophia I had not previously known. It was sent by Mr. H. J.
Vallender who found plants growing near Batang Malaka in Negri Sembilan.
He reported that he had seen the leaves, that they were plicate like those
of Spathoglottis but much narrower, and that the leaves died before the
flowers appeared. I found that the flowers agreed with descriptions cited
by J. J. Smith under E. bicarinata in his enumeration of Sumatran orchids.

Wishing to prepare a description of E. bicarinata for my manuscript
on orchids of the Malay Peninsula, I looked up the descriptions cited by
J. J. Smith (they did not include Pantling’s drawing) to discover details
of stem and leaf, but found no such information. The original description
of E. venosa (one of the synonyms, based on a Queensland specimen) called
the plant a leafless herb. It was this that led me to write for further
information from Australian botanists. I had seen the rather “ sketchy ”
fine-drawing of E. venosa in Bailey’s Catalogue. This showed no leaf but
it showed a curious prostrate tuber-like rhizome very different from any
other Eulophia I knew. I did not realize that this was an early drawing
by C. T. White, and I questioned its accuracy. Mr. White replied that he
believed it was accurate, but that he had not seen a living plant of the
species for many years.

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

Soon afterwards, Mr. Vallender sent two pieces of resting rhizomo
which were clearly of the same nature as that shown in White’s drawing.
They were whitish, tapered to both ends, and bore ring-shaped bases of
decayed scale-leaves. It appeared that they grew horizontally, and they
were slightly flattened. The remains of former inflorescences were not
clearly distinguishable.

The two pieces of rhizome were planted in pots, and after some weeks
each produced a slender erect shoot which at first consisted only of purplish
sheaths. Later, foliage-leaves appeared from within the sheaths, and their
development is not yet completed. The first leaf on the larger plant grew
to a length of 35 cm. above the protecting sheaths, with a maximum width
of 15 mm. The second leaf grew much longer though no wider, and had a
distinct stalk. Both leaves were so weak that they drooped and later
became folded across the middle.

At this stage I removed the larger plant from the pot and washed it
carefully. The leafy shoot had arisen from a bud in the axil of one of the
scale-leaves on the old rhizome ; the shoot grew straight upwards and had
only a slightly swollen base which bore a few roots. The old rhizome was
slightly shrunken. Directly beneath the erect leaf-shoot, and growing
downwards , was a new fleshy organ, more or less ovoid, with a broadly
rounded apex ; its surface was covered with colourless scale-leaves.

The habit thus appears to be a sympodium consisting of alternate
very unequal elements, contrasting with the sympodium of equal elements
in the other species of Eulophia in Malaya. In E. graminea , for example,
there is a succession of equal ovoid pseudobulbs, each arising from the
base of the previous one, each bearing a few narrow leaves on its upper
part, and each later bearing a few inflorescences from its lower part after
the leaves have withered. In E. bicarinata there appears to be first a
short erect non-pseudobulbous leaf- bearing stem with adventitious roots
at its base, and then a tuberous rhizome (covered with scale-leaves) which
rests after the foliage-leaves have gone and can later bear a new leaf-shoot.
I have not been able to investigate the origin of the tuberous organ ; pre-
sumably it arises in the axil of one of the basal sheaths of the leaf-bearing
shoot. It seems that the tuberous rhizome begins growth downwards and
continues horizontally ; whether it ends in an inflorescence- bearing part is
not clear. There is the further possibility that the rhizome can bear foliage
leaves near its apex, that it normally does so after resting, and that in the
specimen seen by me a lateral bud developed because the terminal leaf-
bearing part had been injured. Until I have had a plant under observation
for some time longer, I cannot give a final answer to these questions. I
write this note however in the hope that some other observers may be
able to study a living plant in its natural habitat and report on its way
of life.

Looking over taxonomic descriptions of many tropical monocotyledons
of various families, I am impressed by the very cursory references to the
subterranean parts. The variety of vegetative habit is one of the most
astonishing features of monocotyledons. It is no doubt connected with
their lack of a cambium, which prevents their assuming the tree-form and
necessitates the development of adventitious roots. The shape of the
successive parts of the stem, and their relation to one another in mono-
cotyledons which have sympodial growth, and especially in those which
form resting or storage organs, is extremely varied. A most remarkable
series of examples are found within the genus Dioscorea as described in

VEGETABLE HABIT IN THE GENUS EULOPHIA.

various works by Burkill. These examples show a great plasticity in
vegetative form, the ability to change readily in adaptation to new conditions
of growth owing to climatic or other habitat differences. We see
this plasticity also in the genus Eulophia, though on a much smaller scale
than in Dioscorea.

There are doubtless many similar curious facts hidden under such
statements as “ rootstock tuberous,” or “ roots fibrous ” in our older
taxonomic works, and the student of tropical plants may find much of
interest in the investigation of plants so described.

The wide geographic distribution of most of the Indo-Malaysian species
of Eulophia is no doubt to be explained by the fact that they are plants of
open places, not of moist shady forest. The fact too that they are mostly
seasonal in growth and flowering makes them unsuited to the very uniform
climate of Malaya in which dry seasons are neither long enough nor regular
enough to induce regular periods of rest. Many such widely distributed
species of seasonal habit do not occur in southern Malaya though it falls
within their geographic range.

I conclude with a statement of the synonymy of Eulophia bicar inata,
as given by J. J. Smith, with the additional references to illustrations.

Eulophia Mcarinata (Lindl.) Hook. fil. FI. Brit. Ind. 6 : 6 (1890) ; King
& Pantl. in Ann. R. Bot. Gard. Calc. 8 : 180, t. 244 (1898).

Cyrtopera bicarinata Lindl. Gen. et Sp. Orch. 190 (1833).

Eulophia venosa Rchb. fil. in Benth. FI. Austr. 6 : 300 (1873) ; Bailey
Compreh. Cat. Queensl. PI. 529, fig. 515 bis (1913).

Cyrtopera papuana Kranzl. in Notizbl. Bot. Gart. Berlin 2 : 104 (1898).

Eulophia venosa var. papuana Schltr. Orch. D. N. Guin. 416 tab. 145>
no. 545 (1912).

E. versteegii J.J.S. in Bull. Dep. Ag. Ind. Neerl. 19 : 24 (1908) ; Nova
Guinea 8 : 25, t. 9, 5 (1908).

E. neo-pommeranica J.J.S. in Nova Guinea l.c., 26.

Cyrtopodium parkinsonii F. Muell. et Kranzl. in Oesterr. Bot. Zeitschr.
44 : 256 (1894).

Vol. LXIL, No. 6.

RHEOPHYTES.

By C. G. G. J. van Steenis, Rijksherbarium, Leiden, Holland.
(With Plates II. and III.)

(Issued separately, 15th August, 1952.)

SUMMARY.

Rheophytes are plants confined to stream-beds and, though they belong to widely
different families, they have certain morphological characters in common. A preliminary
list of those occurring in Australia is given, together with an amended account of the
genus Torrenticola.

Rheophytes are, by definition (van Steenis 1932), plants distributionally
confined to stream-beds, below flood level. They occur especially in swift
running water of streams subject to overflowing.

Plants of very different structure and systematic status belong to
this class of plants which has hitherto been little recognised as a separate
ecological group. This is rather strange because streams are generally, in
the tropics, the natural highways of the forest.

To those who are acquainted with some typical rheophytes, it is clear
that they deserve to be classed together. Rheophytes occupy a special
ecological niche in nature. As 1 observed several rheophytes in Australian
waters, I want to focus the attention of Australian botanists on this
specialized group, with which I first became acquainted in my boyhood
at Utrecht, Holland. In a swift-running stream near this town, the identity
of a gregariously occurring, submerged, ribbon-leaved water-plant which
never flowered puzzled me for a long time. In passing, it appeared to
represent a rheophytous variety of Sagittaria sagittifolia known exclusively
from streams and rivers.

Since that time I have paid special attention to plants restricted to
stream-beds and have found some similarity in their behaviour. I have
also gradually accumulated a large number of remarks and notes in literature
from all parts of the world and extracted data from floras, plant lists, and
expedition reports. In all, I have notes on several hundred rheophytes, and
it appears that they occur all over the world both in wet and semi-arid
countries.

H. N. Ridley (1893) in a report on the flora of the east coast of the
Malay Peninsula wrote :

“ The hills which enclose the rocks are upward of 2,000 feet in altitude, and are
composed of quartzite, clad to the top with tall trees. One could not but be struck
with the similarity in form of the foliage of the chief plants which grew close to the
water’s edge. There was here quite a series of plants with peculiarly long, narrow,
acuminate leaves, such as the specific name salicifolia suggests. They belonged to
all manner of orders and included a Galophyllum, an Ixora (I. stenophylla Miq.), a
Hygrophila ( H . saxatilis n.sp.), an Antidesma (A. salicifolia Miq.), a Didymocarpus
( D . salicina n.sp.), a new genus of the Asclepiadaceae ( Spiladocorys ), a Podochilus
( P . maingayi Hook, f.), a Ficus ( F . pyriformis Hook.), a Rhynchopyle, and a Melastoma.

The forms of leaves in the tropical woods bear a great relation to the amount and
force of the rainfall. During a tropical shower it is easy to see how well the commonest
form of jungle leaf (a lanceolate, acuminate or caudate one) is adapted for throwing
off the heavy rain rapidly. But the form of the narrow leaf among the plants mentioned
above seemed to me to be the result of adaptation to avoid another danger.

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

All these plants grow on rocks near the water’s edge, and as the river is subject to
very rapid rises from sudden falls in the mountain districts, these plants are liable to
submersion in a violent rush of water. Broad foliage would be torn off or mutilated,
but narrower leaves, offering a less resistance to the water, would receive no injury.
One may compare with this the narrow floating leaves of the aquatic Ranunculi and
Potamogetons which inhabit the rapid streams in Europe.

The islands of the Tahan river formed of boulders produced several interesting
plants, including Homonoia, not hitherto recorded south of Siam , a curious new and
shrubby Phyllanthus, and abundance of the beautiful Arundina speciosa Bl.”

This excellent description was preceded by observations of the great
Italian explorer O. Beccari whose report, however, appeared later and is
hidden in the pages of that most interesting book “ Nelle foreste di Borneo ”
with the English translation “ Wanderings in the great forests of Borneo ”
(1904). On page 392 of the latter he wrote : —

“ Stenophyllous plants. I have adopted this term for certain plants growing on
river banks or in the beds of torrents, which have linear or else very narrow leaves —
narrower than those of congeneric species growing in the forest. By this I do not mean
that every plant with narrow leaves must of necessity grow by the side of rivers nor
that it is only in Borneoan rivers that such are observable. That plants with linear
leaves have such a habitat is well known, and I need only mention as an example the
many species of Salix so frequent in such localities.

In Borneo, however, along the inland rivers, stenophyllism appears to me much
more accentuated, and more instructive by reason of the fact that the plants exhibiting
this characteristic are numerous, and in many cases very strictly localized. They thus
give one the idea of growing on the spot where they modified their structure, and more
©specially their leaves, in compliance with the stimulus received. I am inclined to
ascribe the stenophyllism as due to the action of the continuous currents of air, so
constant along rivers, and secondly, to that of periodical floods. In the latter case
stenophyllism is associated with great flexibility and toughness of the stems and
branches, such as that exhibited by several species of Salix (here follow many
examples of plants collected by him). ... I believe also that, although these species
appear to be highly localized forms, they may be found along other rapid rivers subject
to sudden inundations.”

The observations of Beccari eminently supplement those of Ridley and
leave no doubt about the existence of rheophytes. Many later collectors
and writers have recognised them in collections, e.g., Merrill, in his studies
on the Philippine and Bornean flora, and Endert, Beumee, Burkill, Holttum
and Corner.

Beccari rightly drew attention to the fact that it is not strange that
these plants are suited to stream- bed conditions, but that the main question
is : why are they, in nature, limited to the part below flood-level ?

As in similar ecological groups, such as the saprophytes, hydrophytes,
myrmecophytes, cushion plants, epiphytes, ephemerals, geophytes, etc.,
the representatives show a number of parallel adaptive characters in common
but their representatives are scattered through all orders of the vegetable
kingdom. Rheophytes are known among the Cryptogams (Algae, Bryo-
phytes, Pteridophytes), and among the lower organisms they are mostly
known as torrenticolous forms. In zoology the term rheobiology has been
coined for studies of organisms confined to running waters. Among the
Pteridophytes, which comprise a rather large number of rheophytes, the
most singular representative is the leafless Ophioglossum inconspicuum
v.A.v.R. occurring in Papua in gravel-beds covered with Saccharum
spontaneum. According to my numerous data, rheophytes occur in about
60 families of flowering plants. Some families contain a large number,
e.g., the Acanthaceae, Araceae, Moraceae (mainly Ficus), Myrtaceae,
Euphorbiaceae, Compositae, and Rubiaceae. Most unexpected families and
genera have produced rheophytous species, e.g., & Quercus in Hainan, Buxus in
the Philippines, and Notelaea in New Caledonia.

RHEOPHYTES.

Very often rheophytes can be recognised in floras or plant lists by
their specific epithet referring either to habit or leaf shape which has struck
the collector or the phytographer ; epithets like salicinus, rheophila,
stenophylla, riparia, lanceolata, viminea, virgata, linearis, and salicoides have
often been given to rheophytes.

Among rheophytes there is only one family of which all members are
confined to rapids and torrents, viz., the highly specialised Podostemaceae.
An emended description and a plate of the only Australian representative
of the family are given below.

Of the other families, there are some genera, often monotypic, which
consist exclusively of rheophytous species, e.g., Rhabdia (Borraginaceae),
Nerium (Apocynaceae), Enaulophyton (Melastomaceae), Myrmeconaudea
(Rubiaceae), Fentasacme, and Dorystephania (Asclepiadaceae).

Most frequently, however, a genus includes only one or a few
Theophytous species, but more than one occur in Neonauclea (Rubiaceae),
Antidesma (Euphorbiaceae), Ficus (Moraceae), Syzygium ( Eugenia )
(Myrtaceae), Aglaia (Meliaceae), etc. Rheophytes may also be represented
by a subspecies or a mere variety of some non-rheophytous species. Among
Gramineae, Cyperaceae and Liliaceae, several species belong to the
rheophytes ; their representatives are naturally stenophyllous.

Also among water-plants (hydrophytes) some species are typical
rheophytes. In a certain sense all rheophytes are hydrophytes, but it
should be realized that practically all are perennial, and that the majority
are woody, suffruticose or shrubby, 1-2 m. tall, or even small to medium-
sized trees. Some of the latter are, hence, only typical rheophytes (subject
to the current) in their juvenile stage ; in their later stage their crown is
elevated above the stream and no longer affected by floods. Anyhow, the
woody rheophytes have been derived from land-plants, behave as such, and
at least find their closest relatives among land-plants. Moreover, most
hydrophytes possess hygrophytic leaves, but the foliage of rheophytes is
definitely not hygrophytic, but rather tough and firm, subcoriaceous to
coriaceous in the herbarium. None of them has membranaceous leaves.
Doubtless their leaves are provided with fibres or other sclerenchyma.
Probably rheophytes are more heliophilous than hygrophilous. Hardly
any rheophyte possesses hairy leaves, and hardly any has toothed leaves ;
if present, marginal teeth and incisions tend to be of the crenate type.
When cultivated, the structure of the leaves proves to be absolutely constant,
as I have observed in Nerium, Homonoia, Eupatorium riparium and others.

The stems of the rheophytous trees are sometimes distinctly stream-
lined in section, possibly owing to tensions in the stem and the symmetrical
root-system which is elongated in both ways in the direction of the stream.
Also, the crown is oblique, the water- current having a similar distorting
effect as constant winds can have on trees and shrubs (wind-forms).

The equipment of the rheophytes shows three specializations in con-
nection with the habitat, namely : —

( a ) Strong and wide root- system able to anchor in moving gravel

and boulders, or very thick root-matting to adhere to the
rocks.

(b) Tough, flexible structure of stems because of abundance of

fibres in order to give resistance to the swift current in the
struggle for self-preservation and maintenance.

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

seeding, germination of seeds, and dispersal methods of seeds.
Seeds of Podostemaceae are able to stick to surfaces. In many
other rheophytes seeds or fruits are able to drift.

In habit, many shrubby rheophytes have some typical field- characters

in common.

(d) Stream-lined leaves, very often more or less falcate and

attenuate, with high leaf-index, i.e., length divided by width,
exceptionally 2 (in Myrmeconauclea), mostly much more than
3 ; that is, they are at least lanceolate.

( e ) Broomlike, umbrella- shaped or table-shaped habit of the

crown, caused mostly by sympodial, often virgate branching
in such a way that the foliage comes more or less in one
plane. The crown is often oblique and elongated in the
direction of the stream.

(/) Reductions frequently occur in genera with compound leaves,
e.g., in the Pteridophytes and Meliaceae, where the leaves of
the rheophytic members of a genus are reduced by comparison
with allied species, sometimes simple.

Once one has recognised rheophytes, these plants can be picked out
of a collection at sight (with the exception of Monocotyledons) and field
notes on the label generally confirm one’s impression, as I have found on
many occasions. Next to the $afo’;r-like leaves, they have something peculiar
in their habit difficult to define in words. To call them “ virgate ” is too
strong : specimens have often mud or drift between the foliage left after
flooding, and sometimes they have been slightly damaged by the flood
waters.

A noteworthy experience is that rheophyte communities hardly ever
consist of one species : mostly several are found together in colonies where
the current, and hence the “ soil,” is suitable. Rivers and streams are the
gutters of Nature where, among the debris, diaspores of plants belonging
to the most different ecological niches accumulate.

To find out whether a plant can be defined as an obligatory rheophyte,
expert field -knowledge is always needed, and sometimes it can not be
judged locally. One has to look up as many specimens in herbaria as
possible, and check literature. Unfortunately most handlists and floras
seldom define the ecological habitat of the species. Also, collectors often
forget to make a concise note on the habitat of their specimens for the
herbarium. In checking whether a species found in a river-bed below
flood level is a rheophyte, one ought to make a study of its geographical
distribution. Most plants from river-beds are occasionals ; they are often
able to germinate and grow in the river-bed, but their station is mostly
temporary, and depends on regular and fresh supplies of diaspores by
which their stations may appear continuous. Sometimes hydrophytes,
saxophytes or hygrophytes may be able to find a sheltered place on rocks
or under rocks where they can survive locally because they are not exposed
to the direct force of the stream. These facultative rheophytes representing
the bulk of the stream-bed flora, can easily be distinguished on the strength
of their occurrence elsewhere outside the rheophytic habitat.

RHEOPHYTES.

In Australia, some difficulties are encountered, because many water-
courses have, specially in dry regions, accumulations of a varied flora ; also
the height of the floods is very variable, and many streams are typical
“ wadis,” that is, they are flooded at long intervals. Here, one ought to
be careful not to confuse riverine species — that is, those which occur along
streams or in well- watered ravine-beds — with true rheophytes.

If one wants to look for rheophytes, attention should be focussed, not
on sluggish, silted stream-beds, but on streams containing clear water with a
bottom consisting of coarse sand, pebbles, boulders or rocks and streams
with cascades and rapids. An examination should be made of gravel, bars,
and crevices of rocks and ledges.

During my rather hurried visit to East Australia (July- August 1950) I
got the impression that Australia harbours many rheophytes, some of which
I observed through the kind help received from all colleagues whom I
met, and many hints from memory or literature were given.

My Australian records have not been properly checked with herbarium
collections or with the literature, some of which is not available to me.
Therefore, this provisional list is more an invitation to criticism and a
request for additional information than anything else. I will be extremely
grateful for help in tracing true rheophytes. Some time I intend to arrange
my growing number of notes into book form.

Casuarinaceae :

Casuarina cunninghamiana Miq. ! River oak. Seen near Canberra.

Compositae :

Olearia argyrophylla F. Muell.

Cyperaceae :

Car ex alsopkila F. Muell.

Car ex gaudichaudiana Kth.

Car ex polyantha F. Muell.

Leguminosae :

Acacia salicina Lindl.

A. stenophylla A. Cunn.

Liliaceae :

Lomandra longif olia Labill. vel. aff. An apparently distinct
rheophytic variety or subspecies or an allied species was seen near Brisbane.

Malvaceae :

Plagianthus pulchellus A Gray.

Myrtaceae :

Callistemon salignus Sweet.

C. subulatus Cheel.

C. viminalis (Sol.) Cheel. Red bottle-brush. Seen near Brisbane and
Canberra.

Eucalyptus camaldulensis Dehnh.

E. coolabah Blakely & Jacobs.

E. largiflorens F. Muell. (syn. E. bicolor A. Cunn.) should be a
rheophyte in the river Murray and tributaries.

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

Eugenia ( Syzygium ) eucalyptoides F. Muell.

E. smithii Poir.

E. tierneyana F. Muell.

E. ventenatii Benth. Weeping myrtle. Seen near Brisbane.

Melaleuca bracteata F. Muell. River tea-tree.

M. leucadendron L. var. mimosoides, var. saligna, and var. argentea
according to Mr. White. Weeping tea- tree.

Tristania laurina R.Br. Water gum. Seen near Brisbane.

T. neriifolia R.Br. Water gum.

Podostemaceae:

Torrenticola queenslandica Domin.

Polypodiaceae:

Lomaria discolor Mett. (probably hygrophyte ?)

L. lanceolata Sturm. do. ?

L. fluviatilis Mett. do. ?

Proteaceae:

Lomatia myricoides Gaertn. ! (syn. L. longifolia R.Br.). Apparently
one form only. Seen near Canberra and Sydney.

Thy melaeaceae :

Pimelea axi flora F. Muell.

Violaceae:

Hymenanthera angustifolia R.Br. (syn. H. dentata R.Br.).

A note should be added about the use made of some rheophytes, for
with the exception of the Podostemaceae, a number of them can be cultivated
outside streams. Ornamentals are Gardenia jasminoides Ellis and Nerium
oleander L. For Nerium, I checked the habitat of all wild specimens in the
Mediterranean and Near East in the Kew Herbarium, and it proved to be
a true rheophyte. The gregarious growth and ability to form extensive
root systems have induced man to use rheophytes for protection of the
soil against erosion. I found Homonoia riparia planted in North Sumatra
along and in watercourses between steep rice-fields for protecting the
banks. For protecting steep road-talus along roads in tea and chinchona
plantations in West Java, Eupatorium riparium and E. triplinerve are exten-
sively used. The tough, flexible stems have induced man to use Salix in Europe
and elsewhere for matting purposes. This finds its equivalent with the
Dayak people in West Borneo who use Ardisia linearifolia Miq. for making
traps and baskets for fishing purposes.

Emended description of Torrenticola queenslandica Domin.

The oldest record of a Podostemacea occurring in Queensland is the
note by F. Mueller (1882) where he mentions a “ genus and species not yet
determined,” a fact recognised by F. M. Bailey (1901) who added “ that the
authority rested upon specimens collected by Mr. Walter Hill at the John-
stone river in 1873. In 1874, Oliver sent specimens to E. Weddell who
recognised the species as new, but could not place it.” Domin studied this
material and described it provisionally as representing a new monotypic
genus, Torrenticola queenslandica with the alternative name Podostemon
torrenticolum (Damin 1925). His description was based on fruiting material
only.

RHEOPHYTES.

In 1935 the late orchidologist C. E. Carr collected in the Laloki river
cataracts, near Rona, Papua, the same species in fruit and flower. In 1947
I identified this with the Queensland plant of Domin, and later re-
produced an ample figure (1949). In July 1950 I discussed this neglected
plant with Mr. C. T. White with whom I had the great privilege of staying
seven memorable days at Brisbane. Mr. White, who showed me unforgett-
able kindness and friendliness, gave me a little tube containing some
sterile shoots of a tiny plant which he correctly took to be Torrenticola
queenslandica, but most probably representing a vegetative growth different
from the fertile stems. This difference of sterile and fertile shoots has been
observed in many Podostemaceae. The material was collected by Mrs.
M. J. Mackerras, Oct. 12, 1949, in Babinda Creek, near Cairns, NE. Queens-
land, growing on smooth granite boulders in fast water. This is a second
locality for Queensland, and more records can be expected. Fertile material
can be collected during the dry period when the level of the water is low.

The description of the species, copied from the Flora Malesiana, is as
follows : —

Torrenticola.

Domin, nom. prov. Bibl. Bot. 89, 2 (vol. 20) (1925) 149, t. 35, f. 7-13 ;
Engl, in E. & P. ed. 2, 18a (1935) 484 ; Steen. J. Arn. Arb. 28 (1947)
421 ; FI. Mai. 42 (1949) 66. — Podost. indet. F. v. M. Syst. Cens. (1882)
23 ; 2nd Cens. (1889) 166 ; Bail. Compr. Cat. (1913) 417.— Podos-
temon sp. Domin, l.c., nom. alt.

Roots Ungulate, sparsely ramified ; sprouts very close, erect, mostly
simple, rarely branched, thin, ridged, densely foliated. Leaves distichous,
equitant, base laterally compressed, obtriangular, upper ones toothed,
minute, thick, 1-2 carinate, decurrent, lowest semi-amplexicaulous, blade
d: patent. In sterile submerged shoots the central tooth of the leaves is
elongated into a fihform appendage. Flowers single, apical, strongly
asymmetric. Spathella sub-obhque, tipped, irregularly circumscissile-
dehiscent. Flower shortly stalked. Tepals 2, narrow, small. Stamen 1,
articulate with the pedicel, in bud appressed to the ovary, filament broad,
anther broad, connective emarginate, cells introrse. Pollen grains 2-celled.
Ovary dz globular, smooth, 2-locular with 2 grooves where the dissepiment
is attached ; stigmas 2, oblong -lanceolate, in bud appressed to the ovary
towards the stamen. Fruit shghtly oblique, terete, the largest valve
persistent, ribs 10, indistinct. Placenta globular, surrounded by a thin
narrow membranous dissepiment, caducous. Seeds numerous sub -angular
oblong.

Distr. Monotypic, Queensland and Malaysia : SE. New Guinea.

Note. Vegetatively, Torrenticola reminds of certain S. American
species of Podostemon , but the flowers and fruit indicate its affinity with
the Malaysian Cladopus.

Torrenticola queenslandica Domin, l.c. ; Engler, l.c. ; Steen 11. cc. —
Podostemon sp. Domin, l.c. nom. altern.

Roots ca -|-2 mm. broad. Stems 2J-6| cm. long, densely set, dark-
green. Leaves ridged on the back, ca J-2 mm. long. Filiform appendages
in sterile specimens to 15 mm. long. Spathella di sessile, to dz i mm.
stalked, d: 2 mm. long. Pedicel J-1J mm. in bud, 2 mm. in fruit. Tepals
dz | mm. long, filament d: i mm. long, thecae dz f mm., cells slightly
unequal, pollen grains dz 30 p. Ovary 1J mm. diam. ; stigmas d: i mm* »
seeds ca J-J mm. diam.

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

Queensland : Johnstone River, W. Hill, 1874 ; Babinda Creek,
Mrs. M. J. Mackerras, Oct. 12, 1949 (Herb. Brisbane).

Papua : Laloki River, near Rona, C. E. Carr 12415, 1935 (Herb.
Brisbane, Canberra, Melbourne).

REFERENCES.

Bailey, F. M., 1901. Queensland Flora 4 : 1277, Brisbane.

Beccabi, O., 1904. Nelle Foreste di Borneo. Firenze, 1902. Translated into English
by Giglioli, E. : Wanderings in the great Forests of Borneo. London,
1904.

Domin, K., 1925. Bibliotheca Botanica 20, heft 89 (2) : 149, t. 35, fig. 7-13.
Mueller, F. von, 1882. Second Census of Australian Plants: 23. Melbourne.

Ridley, H. N., 1893. On the Flora of the Eastern Coast of the Malay Peninsula.
Trans. Linn. Soc. Lond. Bot. ser. II, 3 : 267-408 (see pp. 269-70).

Tisdall, H. T., 1887. A Ramble in a Gippsland Gully. Viet. Nat. 4 : 67-71.

van Steenis, C. G. G. J., 1932. Botanical Results of a Trip to the Anambas and Natoena
Islands. Bull. Jard. Bot. Buitenz. ser. Ill, 12 : 151-211 (see pp. 197-201).

, 1949. In FI. Males. 4 (2) : 67.

White, C. T., 1930. Queensland Vegetation. Handbook for Queensland, Australasian
Association for the Advancement of Science, 53-62 (see p. 61).

Proc. Roy. Soc. Q’land, Yol. LXII., No. 6.

Plate II.

Torrenticola queenslandica Domin. a-b, Branched and unbranched stems ; c, lower
part of stem ; d, central part ; e, upper part ; f-g, stem-bases attached to Ungulate
roots on pieces of gravel ; h, leaf types ; i, bud on apex of stem ; j, closed spathella
with bud ; k, spathella opened artificially ; 1-m, dorsal view of flower ; n-p, larger
valve of fruit ; q, unopened fruit, (a-b, x f , c-q, x 8).— Courtesy Flora Malesiana.

Proc. Roy. Soc. Q’land, Yol. LXII., No. 6.

Plate III.

Torrenticola queenslandica Domin. Specimen from Babinda Creek, stem x 2,
separate leaf x 3.

Vol. LXII., No. 7.

PSEUDORAPHIS SP1NESCENS (R.Br.) n. comb.,
AND SOME RECORDS OF NEW SOUTH

WALES GRASSES.

By Joyce W. Vickery, National Herbarium of New South Wales, Botanic.

Gardens, Sydney.

(Issued separately, 15th August, 1952.)

SUMMARY.

A new combination in the Gramineae is proposed; six grasses are reported from
New South Wales for the first time.

Pseudoraphis spinescens (R.Br.) n. comb.

Panicum spinescens R.Br. Prodr. 193 (1810).

Chamaeraphis spinescens (R.Br.) Poir. Encycl. Meth., Suppl. 2 : 189
(1811).

Chamaeraphis spinosa Beauv. ex Roem. et Schult. Mant. 2 : 253
(1824), apparently based on Panicum spinescens R.Br.

Panicum asperum Koen. in Naturforscher. 23 : 209 (1788) ; not

Panicum asperum Lam. (1778).

Chamaeraphis aspera (Koen.) Nees in Wall. Cat. Herb. Ind. No. 8679
(1849).

Pseudoraphis aspera (Koen.) Pilger in Notizbl. Bot. Gart. Berlin 10 :
210 (1928).

Panicum abortivum R.Br. Prodr. 193 (1810). (See also Kunth Enum
PI. 1 : 131 (1833).

Chamaeraphis abortiva (R.Br.) Poir. Encycl. Meth. Suppl. 2 : 189 (1811).

Orthopogon abortivus (R.Br.) Spreng. Syst. Veg. 1 : 306 (1825).

Chamaeraphis spinescens var. parvispicula Benth. El. Austral. 7 : 499
(1878), based on Panicum abortivum R.Br.

Pseudoraphis abortiva (R.Br.) Pilger in Notizbl. Bot. Gart. Berlin 10 :
210 (1928).

In the Herbarium of the Royal Botanic Gardens, Kew, there are two
sheets from Robert Brown’s collections, No. 6126 named Panicum
spinescens and No. 6127 named P. abortivum. Each sheet bears two speci-
mens, and from the appearance it seems that one piece from each sheet has
been transposed. There is no doubt that these sheets bear parts of the
type- collections of Robert Brown’s two species. On comparison I could
find no significant difference between them. There is a slight difference in
size of the spikelets, but I have found as great a difference in size of spikelets
on the one specimen in various Australian collections. I am therefore
disposed to regard them as conspecific.

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

The species is now referrable to the genus Pseudoraphis Griffith ex
Pilger, and it becomes necessary to select the appropriate specific epithet
available for use in this genus. Pilger referred Panicum spinescens to
Pseudoraphis aspera , based on Panicum asperum Koen. described in 1788
from an Indian and Malayan grass. But this is a later homonym, being
pre-dated by Panicum asperum Lam. (1778). The epithet was subsequently
used in Ghamaeraphis by Nees in 1849, and for priority considerations
would rank only from that date. It is necessary therefore to select one of
the earlier epithets simultaneously used by Robert Brown. Article 56 of
the International Rules provides inter alia that “ If the names or epithets
are of the same date, the author who unites the groups has the right of
choosing one of them. The author who first adopts one of them, definitely
treating another as a synonym or referring it to a subordinate group, must
be followed.” Bentham (1878) appears to have been the first definitely to
select the epithet “ spinescens ” and refer Panicum abortivum to a subordin-
ate group, to Chamaeraphis spinescens var. parvispicula. Stapf (1906, p.
348) also clearly adopted “spinescens” and relegated “abortivum” to
synonymy. The epithet “ spinescens ” must therefore be adopted also
in Pseudoraphis.

In describing Panicum abortivum , Brown refers to Andropogon
squarrosus Herb. Linn. He did not refer to Andropogon squarrosus Linn,
f. Suppl. PI. 433 (1781). It seems evident therefore that he was identifying
a specimen in the Linnean herbarium with his P. abortivum , but was
deliberately refraining from including Andropogon squarrosus Linn. f. in
the synonymy. The identity of Andropogon squarrosus Linn. f. has puzzled
generations of systematic botanists, but only those aspects of the problem
which relate to Panicum abortivum need concern us here. Certain elements
of its description strongly suggest an aquatic member of the Andropogoneae
such as a Vetiveria sp., e.g. “ Flos hermaphroditus sessilis, masculi
pedicellati.” For this reason it has been held by some to apply to Vetiveria
zizanioides (L.) Nash (syn. : Andropogon muricatus Retz.), although this
view was not held by Stapf (1906, p. 346). Moreover, the description of the
glumes could apply to Panicum abortivum only on the assumption that
Linnaeus failed to observe the short lower glume. There seems no doubt
that there is a specimen in the Linnean herbarium which agrees with P.
abortivum ; Brown’s observations on this specimen were confirmed by
Stapf (1906, p. 348). However, the problem remains as to whether this
specimen is in fact that from which Linnaeus drew up his description of
Andropogon squarrosus, and therefore whether it is the type of the species.
In view of the confusion which already surrounds this name, it seems
undesirable to adopt it in Pseudoraphis until the matter has been placed
beyond all doubt.

Chase (1925, p. 203) has taken a different view, and adopted the epithet
“ squarrosus ” for Panicum abortivum and probably P. spinescens , treating
Brown’s reference to Herb. Linn, as indicating definite synonymy. For
completeness the synonymy involved by this interpretation of the basonym
is given below, but as already indicated this view is not accepted by the
present writer, and these synonyms are not held to be applicable to
Pseudoraphis spinescens.

Andropogon squarrosus L.f. Suppl. PL 433 (1781).

Panicum squarrosum (L.f.) Lam. Encycl. Meth. 4 : 743 (1798).

Anatherum squarrosum (L.f.) Beauv. Agrost. 128, 150 (1812).

PSEUDORAPHIS SPINESCENS AND SOME NEW SOUTH WALES GRASSES. 71

Echinochloa squarrosa (L.f.) Roem. et Schult. Syst. Veg. 2 : 479 (1817).

Orthopogon squarrosus (L.f.) Spreng. Syst. Veg. 1 : 307 (1825).

Chamaeraphis squarrosa (L.f.) Chase in Contrib. U. S. Nat. Herb. 24 :
203 (1925).

Pseudoraphis squarrosa (L.f.) Chase in Jonrn. Arn. Arb. 20 : 313 (1939)-

Pseudoraphis spinescens ranges from India, Ceylon, South China,
Romeo, the Philippines and New Guinea to Australia. It is characteristically
rooted in mud with the stems rising above shallow water in pools or rivers.
It occurs in all Australian States except Tasmania. In New South Wales
it is more commonly found along inland rivers and lagoons, though obtained
by Robert Brown from the Port Jackson district.

Cleistochloa subjuncea C. E. Hubbard in Hook. Ic. PI. 33 : t. 3209 (1935).

This species is known from many collections in Queensland, but only
recently it was obtained in New South Wales, at Gordon Vale, south of
Yelarbon, Queensland, 25-11-1949, F. Hely (NSW 13683).

DimorphocMoa rigida S. T. Blake in Univ. Queensl. Papers, Dept. Biol. 1,
No. 19 : 2 (1941).

Originally known only from a somewhat restricted area on sandstone
ridges and hills in the Leichhardt, Warrego and Darling Downs districts of
Queensland, Blake has informed me that he has since found it at Mt. Maroon,
thus extending its range about 200 miles to the south-east. It is now
recorded from two localities a little west of the Sydney district in New
South Wales, some 600 miles to the south. Here, as in Queensland, it was
found on sandstone ridges. N. S. Wales : Bent’s Basin, Nepean River, 9 —
1913, E. Cheel and J. L. Boorman , (NSW 13688) ; on ridge above
Upper Colo, 29-3-1950, J. Vickery (NSW 11006).

Sehima nervosum (Rottl.) Stapf in Prain, FI. Trop. Afr. 9 : 36 (1917).
(Syn. : Ischaemum laxum R.Br.).

This species extends from Tropical Africa and Tropical Asia to Tropical
Australia. It was first recorded from New South Wales by F. Turner
(1891, 1895), who reported a specimen from Tamworth. It was therefore
accepted as a component of the New South Wales indigenous flora by C.
Moore (1903, p. 85) and by Maiden and Betche (1916). Turner’s collections
are apparently no longer extant, and there have been no specimens of
Sehima nervosum from this State in the New South Wales National
Herbarium until recently. Turner’s record has now been confirmed by a
specimen received from northern New South Wales, some 20 miles north
of Warialda at “ Freestone,” Coolatai, on a stony basalt hill, badly eroded,
with very shallow soil between stones, 29-12-1950, A. Johnson (NSW
13681).

Brachypodium distachyon (L.) Beauv. Agrost. 101, 155, 156 (1812).

This small annual, a native of the Mediterranean region, has been
known in the southern Australian States for many years. As might be
expected, it appears to be advancing northwards into New South Wales,
where its presence is definitely established by the following record : Howlong
to Brocklesby Road, 30-11-1949, E. J. McBarron 4189 (NSW 13690).

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

Phalaris angusta Nees ex Trin. Gram. Icon. 1 : t. 78 (1827).

This native of southern America has been cultivated experimentally
by the New South Wales Department of Agriculture. It now appears to
have become naturalised in at least one area — Windsor district, 12-1949,
J. N. Whitest ( NSW 10592).

Digitaria orbata Hughes in Kew Bull. 1923 : 312 (1923).

This species was described from Herbert’s Creek, Queensland. The
type is in the Herbarium of the Royal Botanic Gardens, Kew, and a specimen
from Cairns in the New South Wales National Herbarium is an excellent
match for it. Some specimens recently collected in northern New South
Wales just south of the Queensland border are referred to this species. It
differs from the related D. diminuta Hughes, a not uncommon species in
New South Wales, in the upper glume which is still further reduced to an
extremely small scale, the back of the upper (fertile) lemma being almost
entirely exposed.

New South Wales : Red Hill, Gordon Vale, south of Yelarbon (Queensl.),
23-11-1949, F. Hely {NSW 10732, 10733, 10734).

REFERENCES.

Bentham, G., 1878. Flora Australiensis 7 : 498-9.

Chase, A., 1925. Contrib. U.S. Nat. Herb. 24 : 203

Maiden, J. H., and Betche, E., 1916. Census of New South Wales Plants, 14.
Moore, C., 1903. Proc. Roy. Soc. N. S. Wales 27 : 85.

Stapf, O., 1906. Kew Bull. 1906 : 346, 348.

Turner, F., 1891. Ag. Gaz. N. S. Wales 2 : 379.

— , 1895. Australian Grasses, 32.

Vol. LXIL, No. 8.

THE SIGNIFICANCE OF THE MALLEE HABIT
IN EUCALYPTUS.

By N. T. Burbidge, C.S.I.R.O., Canberra.

(Issued separately , 29th August , 1952.)

SUMMARY.

The mallee and probably also the marlock form in Eucalyptus are considered to be
of recent origin ; possibly developing in response to the severe change in climatic
conditions which occurred at the end of the Pleistocene. Mallees and marlocks are
found in a number of different sections of the genus but there is no evidence that they
are either primitive or that they have been associated with a particular line of
development. Though they may have had an independent origin in different parts of
the continent, the chief centre of development for the species most conspicuous in the
“ Mallee ” ecological association appears to have been in the Southern Eremaea in
Western Australia. This area is regarded as a secondary centre of speciation for the
genus. The migratory routes of certain birds may have been associated with the
migration of these mallee species to the eastern parts of Australia.

The plant form in Eucalyptus falls into three main categories : (i) Trees,
(ii) Mallees, in which a number of woody stems carrying scanty foliage arise
from an underground woody stock, and (iii) shrubs or mallee-like plants in
which the woody stock is absent or poorly developed. The mallee-like
plants in the last group are commonly referred to as marlocks in Western
Australia and this colloquial term will be used here.

Examples of mallee species are : E. morrisii (Macr anther ae-Exsertae)
from the Flinders Range to western New South Wales ; E. diversifolia
(Renantheroideae- Diver siformae) along the coast from Western Australia to
Victoria ; E. stricta ( Renantheroideae- Fraxinales ), central coast of New
South Wales and the Blue Mountains but not a very typical mallee ;
E. leptophylla ( P or anther oideae-Fruticosae ), Western Australia to Victoria ;
E. gracilis (Gradies- Aridae), Western Australia to New South Wales and
Victoria ; E. cneorifolia (Micrantherae-Eremophilae), Kangaroo Island,
South Australia. The two species most commonly referred to in Australian
literature are E. dumosa (Macr anther ae-Dumosae) and E. oleosa

(Platy anther ae-Subulatae). In both these the circumscription of the species
has been the subject of considerable attention and opinions differ as to the
amount of morphological variation which should be included under the
specific names (Gardner & Watson 1950 ; Burbidge 1947). This makes it
difficult to define the areas of distribution, but if a wide circumscription is
accepted, they extend from the drier areas of southern Western Australia
to similar areas in eastern Australia.

It is evident from the distribution indicated above that mallees occur
in a number of ecological habitats though they are absent from the higher
rainfall zones except where the edaphic conditions are unfavourable for the
development of forest. However, the “ Mallee’’ association, as understood
by ecologists, is characteristic of the drier parts of the winter rainfall belt,
i.e between the wooded and forested areas of the coastal regions and
the arid associations of the dry interior.

A ligno-tuber is found in many seedling Eucalypts and, under
natural conditions, the young plants often have a shrubby form during the
first few years. It is only later that one of the stems becomes predominant

74 PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

and grows out into the main trunk of the sapling. The most frequent
adult form is the tree, but mallees and marlocks occur throughout the
genus and such forms, in which many stems arise from ground level, may
represent the retention of a juvenile state. There is no evidence that either
mallees or marlocks are primitive in Eucalyptus since their chemical and
morphological characteristics are not, at least in the case of the mallees,
those which are usually regarded as primitive by most writers.

The sections of the genus generally considered to be the most primitive
are the Corymbosae and Eudesmieae, though both exhibit a degree of
specialisation which indicates that they cannot be taken as prototypic.
In both groups the essential oils are high in pinene and low in cineol content
(Baker & Smith 1920 ; Watson 1935-36). Leaves of the Bloodwood type
(i.e. Corymbosae ) are amongst the ear best known fossil remains of Eucalyptus
(Cambage 1913). The nature of the essential oils, the leaf-venation, anther-
type and, in some cases, the bark in the Corymbosae as well as the capsule
form in the Eudesmieae show the closest affinities with Angophora. The
Corymbosae nearly all have a tree form but some of the Eudesmieae are
marlocks. Neither group includes any true mallees.

The scattering of mallee species through the genus is demonstrated in
the above list and also in Table I below. In all cases the essential oils are
low in pinene, and the cineol content varies from about 30 per cent, in some
to 90 per cent, or more in the Subulatae (Gardner & Watson 1950). If the
general conclusions drawn by Baker and Smith ( l.c .) regarding the correlation
between increasing oil complexity and development of the genus are accepted,
then it is evident that mallees cannot be considered primitive. This is
further substantiated by their morphology. Though the important group
Dumosae is placed in the same antheral class as the above groups (Blakely
1934) the anthers and pollen grains are smaller and the leaf- venation and
capsule-type show more specialisation and variation in form. In the remain-
ing groups the anthers are specialised but the secondary nature is particularly
shown in the Subulatae where the style is persistent and splits to form the
characteristic fragile points on the valves in the matured fruit. This feature
is not seen in any other Eucalyptus spp. and it has probably appeared
comparatively late in the evolution of the genus.

Consequently it is the opinion of the writer that the mallee form
represents a secondary development in the genus Eucalyptus.

It is possible that the marlocks and other shrubby forms are also of
comparatively recent development, but the case is less clear. Blakely
(1934) does not distinguish between mallees and marlocks so that not all
of his common names can be accepted as indicative of the nature of the
plant in the field, but marlocks are similarly scattered through the genus:
E. eudesmioides , E. tetragona and E. erythrocorys (Macrantherae- Eudesmieae) ;
E. setosa, which may be a low shrub in arid habitats ( Macrantherae -
Corymbosae) ; E. tetraptera ( Macrantherae-Tetrapterae ) ; E. redunca
(Macrantherae- Subcornutae) ; E. kruseana (Macrantherae Globular es) and
E. forrestiana (Platy anther ae-Quadricostatae). All these species except
E. setosa are found in the drier areas of southern Western Australia. The
fact that a number of marlocks are found in the Eudesmieae shows that
secondary deviation cannot be argued on morphological grounds alone, but
it may be significant that all occur in arid habitats as is the case with the
majority of mallees, i.e., they are found under climatic conditions which
are unlikely to have been widespread in Australia during the Miocene
when the major development of the genus probably took place.

Distribution of the More Important Subgeneric Groups of Eucalyptus including Mallee Species (after Blakely).

SIGNIFICANCE OF MALLEE HABIT IN EUCALYPTUS.

Northern

Territory.

Var.

l ^

: ; ; - ; ;

Queensland.

Var.

; : ; - ; ;

Victoria.

Var.

Ph’

Eh'

New South Wales.

Var.

: ^ ^ : : :

—I

T.|

r-H fH r-H LO • •

South Australia.

Var.

Ph*

r-t <M

r-( r-H CO

Spp.

- : : - : :

GO — i lO p-i

Western Australia.

Var.

r-H . • • r-H

E-?

j 2

Spp.

Dumosae

Fruticosae

Aridae

Subulatae

Leptopodae

Contortae

•

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

The question thus arises as to when these smaller forms — mallees and
marlocks — were developed. Conditions during the Pliocene and Pleistocene
were wetter than they are today and forests were more widely distributed
but the pluvial conditions appear to have had an abrupt conclusion if
Crocker and Wood (1947) are correct. According to these authors the
change of climate at the end of the Pleistocene resulted in the destruction
of much of the vegetation of Central Australia and the retreat of the
Australian Element, of which Eucalyptus is so characteristic a constituent,
to the coast, especially towards the south-west and south-east of the
continent. They consider that there was an interim period, following the
onset of aridity, during which biotypes capable of re- colonising the denuded
areas were developed from the plants of the refugial areas.

If the present distribution of the rainfall belts is considered, it will be
seen that the “ Mallee ” association is best developed in the areas between
the zones to which the Pleistocene flora would have retreated and the
more arid interior. Where mallee species, other than those found in the
“ Mallee,” occur, there is always some qualifying feature, e.g., the conditions
peculiar to sand-dunes (E. diversifolia) or the shallow soils of the Hawkesbury
Sandstone (E. stricta ), where the change from the pluvial climate of the
Pleistocene would have been locally marked in its effect on the existing
flora. It is therefore suggested that a large number of the mallees and
marlocks developed as a direct result of the climatic change at the end
of the Pleistocene.

While some of the Subulatae extend into Central Australia, e.g.,
E. oleosa var. glauca (E. transcontinentalis), west of Alice Springs, the paucity
of mallees or marlocks in Northern Australia suggests that these types
cannot have been widespread prior to the arid period. If mallees had
previously existed in the interior then one would expect to find them more
or less encircling the present arid region, i.e., there would have been a
coastward retreat in several directions rather than a southward retreat to
produce the present southern arc. Also there is a lack of representatives in
the refugial areas of Central Australia.

In Table I the distribution of the species most important in the
“ Mallee” association is shown under their various intra-generic groups. It
will be noted that there is evidence that the centre of development lies in
Western Australia and a check of the localities given by Blakely (1934)
shows that they are mainly found in the southern Eremaean Province as
defined by Gardner (1944). It is noteworthy that, whereas the eastern
forms are almost all mallees, the western ones include trees both in the
endemics and in widely distributed forms, e.g., E. oleosa, E. oleosa var. glauca
{E. transcontinentalis), E. flocktoniae and E. gracilis, though the last-named
is seen as a small tree in the Murray Mallee of South Australia. From the
figures there appears to be a secondary centre of speciation for Eucalyptus,
particularly with regard to mallees, in the southern Eremaea, and the eastern
forms with their more pronounced mallee habit may have migrated from
this western centre despite the edaphic barrier of the Nullarbor Plain. If so
the migratory route probably passed to the north of the plain through
country where there are now scattered patches of “ Mallee ” (Wood 1929 ;
Giles 1889).

This view is in direct opposition to that of Crocker and Wood (1947),
since they consider E. oleosa and E. gracilis as having existed in refugia
during the worst of the arid period to expand later into their present

SIGNIFICANCE OF MALLEE HABIT IN EUCALYPTUS.

areas. If they are correct then these species would have retreated to the
outer fringe of the South-Western Province in Western Australia and to
the Flinders Range in South Australia, and the former area could still have
produced the species of the southern Eremaea. This, however, presupposes
that the relics in the Flinders Range, in contrast to the western representa-
tives, retained a minimum amount of inherent variability since there are
no well marked endemics in that area today. This is unlikely as the
Eucalypts of the Flinders Range show considerable variation especially in
the cases of E. oleosa and E. dumosa (Burbidge 1947). A better explanation
than that given by Crocker and Wood is that these mallees originated in
the west and are undergoing further segregation elsewhere. Furthermore
dhe lack of climatically probable refugia and of endemics makes it
unlikely that the mallees of New South Wales and Victoria included in
the groups in Table I are either survivals from an earlier period or the
original stock from which the others arose.

The distribution of the Dumosae strongly supports the above view,
but the inclusion in the Subulatae of E. umbrawarrensis from near Pine
Creek in the Northern Territory and E. pachy calyx from near Cairns in
North Queensland raises certain difficulties. Quite apart from their
distribution however, their morphological characters make it extremely
doubtful if they are correctly placed in the Subulatae . It is more likely
that they represent an offshoot from the same platyantherous stock as
that from which the southern Subulatae arose. E. bakeri in northern New
South Wales and south-eastern Queensland seems to be related closely to
E. oleosa (sens, lat .) and may be a geographic race. The affinities of
E. squamosa , which grows in certain parts of the Hawkesbury Sandstone
in New South Wales, deserve further study. It is platyantherous, but
the capsules and their valves are not particularly typical for the Subulatae.

A recent paper by Gentilli (1949) includes information which appears
to support the idea of a western origin for the mallees dealt with in Table I.
Both the Ring-necked Parrot ( Barnardius zonarius ) and the Sitella ( Neositta
pileata) are presumed to have migrated from a centre in the South-West
Province of Western Australia and to have reached at least the longitude
of the Flinders Range before coming into contact with other races of the
same species. It is inferred that these migrations are recent, i.e., since
the arid period. Similar conclusions with regard to the Sitellas are reached
independently by Mayr (1950). The migration of these birds could have
been contemporaneous with and partly dependent upon the spread of the
mallees, which would have provided nesting and feeding sites. The only
alternative ecological association through which they could have passed
would have been the Mulga ( Acacia aneura), but at present Ring-necked
Parrots are comparatively rare in Mulga areas. The importance of the
Mallee-Mulga boundary in the distribution of birds has been stressed by
Serventy and Whittell (1948) so that a route through the Mallee is more
probable.

REFERENCES.

Baker, R. T. & Smith, H. Gr., 1920. A Research on the Eucalypts and their essential
oils. Sydney.

Blakely, W. F., 1934. A Key to the Eucalypts. Sydney.

Burbidge, N. T., 1947. A Key to the South Australian species of Eucalyptus L’Herit.
Trans. Roy. Soc. S. Aust. 71 (2) : 137-163.

Cambage, R. H., 1913. Presidential Address. Proc. Roy. Soc. N. S. Wales 47 : 18-58.

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

Crocker, R. L. & Wood, J. G., 1947. Some Historical Influences on the Development
of the South Australian Vegetation Communities and their bearing on
Concepts and Classification in Ecology. Trans. Roy. Soc. S. Aust. 71 (1) :
91-136.

Gardner, C. A., 1944. The Vegetation of Western Australia with special reference to
the climate and soils. Jour. Roy. Soc. W. Aust. 28 : xi-lxxxvii (1941-42).

Gardner, C. A. & Watson, E. M., 1950. The Western Australian varieties of
Eucalyptus oleosa F. Muell. ex Miq. and their essential oils. Jour. Roy. Soc.
W. Aust. 34 : 73-86. (1947-48).

Gentllli, J., 1949. Foundations of Australian Bird Geography. The Emu 49 : 85-130.

Giles, E., 1889. Australia Twice Traversed. Vol. II. London.

Mayr, E., 1950. Taxonomic Notes on the genus Neositta. The Emu 49 (4) : 282-291.

Serventy, D. L. & Whittell, H. M., 1948. A handbook to the Birds of Western
Australia. Perth.

Watson, E. M., 1935-36. The Essential Oils of the Western Australian Eucalypts,
Part III. Jour. Roy. Soc. W. Aust. 22 : 113-118.

Wood, J. G., 1929. Floristics and Ecology of the Mallee. Trans. Roy. Soc. S. Aust.
53 : 359-378.

Vol. LXII, No. 9.

OPISTH IOLEPIS, A NEW GENUS OF
PROTEACEAE FROM QUEENSLAND.

By L. S. Smith, Queensland Herbarium, Botanic Gardens, Brisbane.

(With Plate IV.)

(Issued separately, 29th August, 1952.)

SUMMARY.

A new genus of the Proteaceae is described and a new transfer made.

The high degree of endemism in the rain-forest flora of north-eastern
Queensland, despite the predominance of Malaysian genera, has already
been remarked upon by the late C. T. White (1930, p. 234). One peculiar
feature of this flora is that up to the present six monotypic genera
of Proteaceae have been recorded, namely, Austromuellera, Buckingham-
ia, Cardwellia, Carnarvonia , Musgravea, and Placospermum. To these must
now be added Hollandaea and Opisthiolepis. C. T. White, in manuscript,
proposed the following transfer —

Helieia lamingtoniana (F. M. Bail.) C. T. White comb. nov.

Hollandaea Lamingtoniana F. M. Bail, in Queensl. Agric. Journ. 5 : 390
(1899).

The only accepted species of Hollandaea is now H. sayeri F. Muell.
Opisthiolepis , described below, raises the number of genera, with
Queensland representatives, in the tribe Embothrieae to seven.

Opisthiolepis L. S. Smith : genus novum in tribu Embothriearum
ponendum ; inter genera hujus tribus squama hypogyna solitaria bidentata
distinctum.

Flores hermaphroditi, parvi, incur vati, parum irregulares. Perianthii
segmenta 4, inaequilonga, primum a latere abaxiali disjuncta, mox delapsa ;
tubus obliquus, db 4-angulatus, superne vix angustior ; limbus primum
subglobosus, laminis concavis. Stamina 4, ad basin laminarum aflixa ;
filament a complanata, lata, brevia, apicem versus in connectivum latum
expansa ; antherae late ovatae. Squama hypogyna 1, adaxialis, oblonga,
apice bidentata. Ovarium breviter stipitatum, 1-loculare ; ovula ca.
10-12, biseriatim imbricata, oblique disposita, medio vel supra medium
lateraliter aflixa. Stylus compressus, apice versus leviter constrictus deinde
in discum oblique dilatatus, demum decurvatus ; stigma parvum, in medio
disco. Folliculus (pauci delapsi solum visi) recurvus, breviter stipitatus,
demum expansus, sublignosus, subplanus, apice leviter recurvus, bilobatus.
Semina ignota.- — Arbor. Folia alterna, in ramis fertilibus integra, sterilibus
plerumque simplicia pinnataque inter dum sparse dentata. Racemi
axillares, tenues, spiciformes. Flores parvi, brevissime pedicellati, per
paria bractea parva caduca suffulti.

Species 1, Australiae septentrionali-orientalis incola.

0. heterophylla L. S. Smith, species nova. Plate IV.

Arbor mediocris vel alta (usque ad 30 m. sec. Hanson). Ramuli juniores
appresse ferrugineo-tomentosi, leviter angulati vel rotundati, vetustiores
lenticellis parvis pustulatis dense notati, Folia dimorpha, simplicia vel

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

pinnata ; foliorum simplicium lamina coriacea, ovata vel oblongo-elliptica
(4-) 7-15*5 (-30) cm. longa, (1*6-) 2*6-7 (-10*5) cm. lata, apice obtusa vel
breviter acuminata, mucronulata, basi angustata, supra appresse pubescens,
mox glabrata, subtus indumento minuto denso appresso nitente ferrugineo
vel demum d= argenteo vestita, margine integra vel interdum sparse dentata,
nervo medio subtus valde prominente, nervis later alibus prominulis ca.
10-18 utroque latere; petiolus primum appresse tomentosus, ca. 1*2-3 (-5)
cm. longus, supra canaliculatus, subtus rotundatus, basi incrassatus ;
foliorum pinnatorum 28-50 cm. longorum pinnae 5-18, quoad indumentum
nervatioque foliis simplicibus similes, lanceolatae vel oblongo-lanceolatae
(4*4-) 7*6-16*4 cm. longae, 1-9-3-8 cm. latae (vel harum terminalis interdum
major), apice acutae vel acuminatae, mucronulatae, basi inaequilaterales,
margine integrae vel sparse dentatae, petiolulatae petiolulo usque ad 8 mm.
longo decurrente vel interdum superiores sessiles ; petiolus 4*4-7*6 cm.
longus, cum rhachi supra canaliculatus subtus rotundatus. Eacemi axillares,
spiciformes, laxiflori, simplices, solitarii vel interdum geminati, ferrugineo-
tomentosi, 3-9 cm. longi ; bracteae extus tomentosae, intus glabrae, ca.
0*9 mm. longae, emarginatae vel obtusae, caducae. Flores glabri, ca. 2 *5-3*2
mm. longi, brevissime pedieellati pedicellis 0*25-0*6 mm. longis per paria
bractea dz abditis ; perianthii limbus subglobosus, ca. 1 mm. longus, tubus
limbo angustior, segmenta inaequilonga horum lateralia falcata ca. 3 mm.
longa, abaxiale ca. 5 mm. longum, adaxiale ca. 1*7 mm. longum ;
stamina parva, filamentis 0* 1-0*3 mm. longis, antheris ca. 0*5 mm. longis,
0-5-0-6 mm. latis ; squama hypogyna oblonga, 0*8 mm. longa, 0*5 mm.
lata, basi crassior, apice bidentata ; ovarium compressum, ca. 1 mm.
longum, 10-12-ovulatum, stipite ca. 0*3 mm. longo praeditum ; stylus
ca. 2 mm. longus. Folliculus (post dehiscentiam) breviter stipitatus stipite
ca. 6 mm. longo, d= applanatus, 5-6 cm. longus, medio 3-4 cm. latus,
1 *5-2*0 mm. crassus.

Queensland. Cook District : Atherton, C. T. White (sterile), January
1918 (Local name “ Silver Oak ”) ; Atherton, A. G. Hanson 21/1 (type —
flowers), February 1950 (Specimen from a small rain-forest tree, which
attains 100-110 ft. in height and 90 ins. g.b.h. ; pinnate leaves absent on
fertile branchlets) ; Malanda, C. T. White (sterile), January 1918 (Local
name “ Silver Oak”); Millaa Millaa, J. King 10 (sterile), March 1951;
Paronella Park, Mena Creek, ca. 14 miles S. of Innisfail, alt. ca. 50 ft., L. S.
Smith PP. 41 (old fruits) 4 August 1948 (A small slender tree ca. 20 ft.
high near the creek bank ; leaves often both pinnate and simple on the one
branchlet, shiny brown or occasionally silvery beneath). North Kennedy
District : Koolmoon Creek, ca. 11 miles SSE. of Pavenshoe, in rain-forest,
alt. 2,400 ft., L. S. Smith & L. J . Webb 4583 (sterile), 29 September 1950
(A tree 70 ft. high, bole 1J ft. d.b.h., slightly channelled at the base. Bark
3/16- J in. thick ; outer — light brown, marked by fine, broken, slightly
reticulating, longitudinal lines of lenticels ; inner — dark red on the outside,
pinkish brown within with numerous fine, pale cream-coloured radial lines,
when blazed showing a very pale cream or faintly purplish tinged honeycomb -
like network, with brownish flecks, sometimes surrounded by pink, in the
centre of the cells. Sapwood 1J ins. thick, cream for J in., then pale pink
for about 1 in. ; heart wood light pinkish brown) ; Kirrama Range, west
of Kennedy, in rain-forest between Society Flat and Yuccabine Creek,
alt. ca. 1950 ft., L. S. Smith ch L. J. Webb 3200 (sterile), 1 August 1947 (A
small tree 45 ft. high, trunk 7 ins. diam. Outer bark brownish, marked by
short, irregular, longitudinal, lenticellate lines with scattered, short,
horizontal cracks between them, here and there with coarse red-brown

OPISTHIOLEPIS, A NEW GENUS OF PROTEACEAE.

pustules ; inner bark J in. thick, dark red on the outside, reddish brown
within. Wood cream for J in., becoming darker pink inwards. Leaves
simple or pinnate, brownish or greyish beneath).

The genus resembles Lomatia in general structure, but is readily
distinguished by the occurrence of a single hypogynous scale in the flower
instead of three glands. Further, the dehisced follicles of the Queensland
species of Lomatia are almost fiat or very shallowly boat-shaped, often
more or less beaked and not recurved at the apex, much thinner and of a
less woody texture. Other differences may be apparent when the seeds of
Opisthiolepis are known, although the ovules are very similar to those of
Lomatia.

Opisthiolepis heterophylla, especially when young, is quite a showy tree
and worthy of cultivation. The foliage comprises mixed leaf-types, is a
bright green above and shiny brown or silvery beneath. The indumentum
of the leaf-undersurface is somewhat similar to that of Grevillea pinnatifida
F. M. Bail, or occasionally G. hilliana F. Muell. The juvenile or intermediate
leaves, whether simple or pinnate, occasionally bear a few small teeth
which, however, appear to be absent from the simple leaves on fertile
branchlets.

I have seen trees in rain- forests from near Cardwell in the south to
Mossman in the north and growing at altitudes of from approximately
50 ft. to 2,500 ft., though nowhere of very large dimensions. However,
Mr. A. G. Hanson, Forester, Atherton, states that they grow to a height
of 100 ft., the bole attaining 2J ft. in diameter, and therefore of commercial
size.

The small, inconspicuous, almost sessile flowers may account for
flowering specimens not having been collected in the past, and I am much
indebted to Mr. Hanson for diligently searching for and collecting the only
flowering specimens so far obtained.

[. L . S. Smith & L. J. Webb 4853, since collected from Lacey’s Creek,
Mission Beach area, shows in addition that the flowers are white and that
the lower racemes are pendulous, attain 15 cm. in length, and are sometimes
branched. Old fruits similar to those described were obtained from the
ground beneath the tree.]

The generic name is derived from the two Greek words orriaOios ,
hinder, and A ems, a scale, and refers to the adaxial, solitary, hypogynous
scale in the flower.

REFERENCE.

White, C. T., 1930. A new genus of Proteaceae from North Queensland. Kew Bull.

1930, No. 6 : 234-5.

Proc. Roy. Soc. Q’land, Vol. LXII., No. 9.

Plate IY.

5, pistil and hypogynous scale after perianth -segments have fallen x 10 ; 6, hypogynous
scale x 10 ; 7, apex of style with stigma x 10 ; 8, longitudinal section of ovary x 10 ;
9, two ovules after fertilisation x 16 ; 10, tip of lateral perianth segment showing

position of stamen x 10 ; 11, back view of anther from abaxial perianth segment x 10 ;
12, tip of adaxial perianth segment x 10 ; 13, side view of dehisced follicle x 1 ; 14, back
view of dehisced follicle x 1 .

Vol. LXII., No. 10.

THE IDENTIFICATION AND DISTRIBUTION
OF SOME CYPERACEAE AND GRAMINEAE,
CHIEFLY FROM AUSTRALIA.

By S. T. Blake, Queensland Herbarium, Botanic Gardens, Brisbane.

(With Plates V and VI.)

(Issued separately, 29th August, 1952.)

SUMMARY.

The identification, nomenclature and distribution of some Cyperaceae and
Gramineae are discussed. Six species are new records for Australia, one for both
Queensland and South Australia, and two for Queensland, while six new species are
described.

Many sedges and grasses found in Australia are also found over a wide
area beyond Australia. The identification of these “ wides ” is often a
serious difficulty which can only be solved by the study of types of all the
names of allied species, many of which are not readily available. The loan
of several such types from the herbaria at Lund (ld), Paris (p) and Copen-
hagen (c) has been made to me through the kindness of Dr. A. Hassler,
Prof. H. Humbert and Dr. O. Hagerup respectively, and I wish to express
my gratitude to them for their assistance. Miss M. B. Tindale and Mr. E.
Nelmes have kindly copied passages from publications in the library of the
Kew Herbarium which are unavailable in Australia. Some of the species
studied are discussed in this paper ; the opportunity has been taken to
record extensions to the previously known range of some allied endemic
species, to describe some new ones, and to correct an error in nomenclature.

Cyperaceae.

Cyperus suleinux C. B. Clarke in J. Linn. Soc. 21 : 56 (1884).

Queensland. — North Kennedy District : Bowen, near roadside in

granite sand, 25th Sept. 1950, Blake 18544.

New for Australia ; widely spread through Malaysia to India and
Indo-China. The collection cited above consists of a large number of small
plants with culms 1-7 cm. high and the wide- spreading rays of the inflores-
cence up to 5-5 cm. long below the rich brown spikelets. The species belongs
in subgenus Pycreus and its chief distinguishing characters are its annual
habit, very obtuse glumes closely 3 -nerved on the back with nerveless
sides, and oblong nut with a shallow furrow down each face.

Scirpus L. Sect. Actaeogeton Reichb. Ic. FI. Germ, et Helv. 40 (1846) ;

Beetle in Amer. J. Bot. 29 : 653-6 (1942).

This section is based on Scirpus mucronatus L. and its characteristic
features are that the culms are nodeless above the base or have but one
node in the lower part, the leaves are commonly reduced to more or less
mucronate sheaths, the outer involucral bract appears like a continuation
of the culm, and the black or dark-brown or grey-brown nut is marked
by transverse, wavy ridges ; there may be two or three stigmas and
hypogynous bristles may be present or absent ; some species regularly
produce flowers in the leaf-sheaths with greatly elongated styles and stigmas.
Difficulties which had been encountered when studying the Australian
members of this group were not solved by the application of Beetle’s revision

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

cited above. The following discussion is offered as a contribution towards
a better understanding of the species which have been associated in one
way or another with the name S. supinus L. The names and more important
references which must be considered are as follows : —

Eleocharis juncoides (Roxb.) Schult. Mant. 2 : 90 (1824).

Isolepis ambigua Steud. in Zoll. Verz. Ind. Arch, heft 2 : 62 (1854) ;
C. B. Clarke in Philip. J. Sci. 2,C : 99 (1907).

Isolepis ambigua Steud. Syn. Cyp. 91 (1855).

Isolepis (?) juncoides (Roxb.) Miq. FI. Ind. Bat. 3 : 312 (1856).

Isolepis oryzetorum Steud. Syn. Cyp. 96 (1855) (“ oryectorum ”).

Isolepis polycolea (Notaris) Steud. Syn. Cyp. 95 (1855).

Isolepis proxima Steud. Syn. Cyp. 95 (1855).

Isolepis supina (L.) R. Br. Prodr. 221 (1810) ; Nees in Wight Contrib.
Bot. Ind. 108 (1834).

Isolepis uninodis Delile FI. Egypt. 8, t. 6, fig. 1 (1812).

Scirpus cernuus Vahl Enum. 2 : 245 (1806) ; Bedevian, Illustrated

Polyglottic Dictionary of Plant Names 535 (1936).

Scirpus dissachanthus S. T. Blake in Viet. Nat. 63 : 116 (1946).
Scirpus erectogracilis Hayata Ic. PI. Formos. 6 : 114 (1916).

Scirpus erectus Poir. Encycl. 6 : 761 (1804) ; C. B. Clarke in Hook.f. FI-
Brit. Ind. 6 : 656 (1894) ; Cherm. in Arch. Botanique 4 (7) : 26
(1931) ; Beetle in Amer. J. Bot. 29 : 654 (1942).

Scirpus Jnallii Gray Addenda Man. ed. 2 (1863).

Scirpus junciformis Nees in Wight Contrib. Bot. Ind. 112 (1834); nec
Retz. (1791) nec (H. B. K.) Poir. (1817).

Scirpus juncoides Roxb. Hort. Bengal. 81 (1814), nomen nudum ; FI.
Ind. 1 : 228 (1820), ed. Carey 1 : 216 (1832) ; non Willd. ex Kunth
(1837).

Scirpus lateralis Forsk. FI. Aegypt.-Arab. 15 (1775) ; Vahl Enum. 2 : 280
(1806) ; Spreng. Syst. Veg. 1 : 207 (1825) ; C. B. Clarke in Prain FI.
Trop. Afr. 8 : 453 (1902), in syn. ; Christens, in Dansk Bot. Arkiv 4 :
12 (1922).

Scirpus lateralis Retz. Obs. 4 : 12 (1786) ; Nees in Wight Contrib. Bot.
Ind. 108 (1834), in syn. ; C. E. C. Fischer in Kew Bull. 1932 : 70
(1932).

Scirpus lateriflorus Gmel. Syst. 127 (1791).

Scirpus luzonensis Presl Rel. Haenk. 1 : 193 (1828).

Scirpus oiyzetorum (Steud.) Ohwi in Mem. Coll. Sci. Kyoto Imp. Univ.
ser. B, 18 (1) : ? (1944).

Scirpus poly coleus Notaris in Ann. Sci. Nat. Ill, 9 : 326 (1848).

Scirpus supinus L. Sp. PI. 49 (1753) ; Benth. FI. Austral. 7 : 330 (1878) ;
C. E. C. Fischer in Kew Bull. 1932 : 70 (1932).

Scirpus supinus L. var. digynus Boeck. in Linnaea 38 : 700 (1870).
Scirpus supinus L. var. digynus Boiss. FI. Orient. 5 : 380 (1884).

Scirpus supinus L. var. elatior Boeck. in Linnaea 38 : 700 (1870).
Scirpus supinus L. var. hallii (Gray) Gray Man. Bot. N. U.S. ed. 5, 563
(1867).

CYPERACEAE AND GRAMINEAE.

Scirpus supinus L. var. uninodis (Delile) Asch. & Schweinf. 111. FI.
d’Egypte 157 (1887) ; Christens, in Dansk Bot. Arkiv 4 : 12 (1922).

Scirpus supinus L. var. uninodis (Delile) C. B. Clarke in Hook.f. FI.
Brit. Ind. 8 : 656 (1894) et in Prain FI. Trop. Afr. 8 : 453 (1902).

Scirpus timorensis Kunth Enum. 2 : 162 (1837).

Scirpus uninodis (Delile) Beetle in Amer. J. Bot. 29 : 656 (1942) (author
wrongly cited), in Amer. Midi. Nat. 34 : 734 (1945) ; S. T. Blake in
Viet. Nat. 63 : 119 (1946) (author wrongly cited).

Scirpus uninodis (Delile) “ Boiss.” var. hallii (Gray) Beetle in Amer. J.
Bot. 29 : 656 (1942).

The earliest of these names is S. supinus, which was based on European
-specimens. Most of the other names were proposed for plants from other
parts of the world, some of which were originally carefully distinguished
from S. supinus, but most of which have been considered by one botanist
or another to be conspecific with S. supinus or only varietally distinct
from it.

S. lateralis Forsk. was described from a plant collected by Forskahl
in Arabia, to the north-west of Aden. The specimen was lost prior to 1806
(Vahl, 1806) and has not been found since (Christensen, 1922 ; Hagerup,
in litt.). Sprengel (1825) thought that Forskahl’s plant was conspecific
with the later described Isolepis uninodis Delile ; at one time, C. B. Clarke
(1902) followed this opinion and Christensen (1922) was inclined to agree.
Otherwise the species has been usually ignored or placed among the doubtful
species. Forskahl’s description is brief and does not mention flower or
fruit. So far as it goes, it could apply to specimens of Isolepis uninodis
except for the longer and apparently more numerous rays of the inflorescence
and the leafy base ; also it would apply to Scirpus litoralis Schrad. (the
only other species which I have found recorded from Arabia) except that
the height ££ pedalis & saepe cubitalis ” is much less than is usual for the
species. The phrase ££ basi foliosa ” does not very well apply to either
species unless sterile culms are meant, but S. litoralis does sometimes
produce conspicuous basal leaves. The Arabic name quoted by Forskahl
and transliterated by him “ hallal ” (better, ££ hhalal ”) is applied
to S. cernuus in Egypt according [to Bedevian (1936), but Forskahl’s
description certainly does not apply to this species. From the available
evidence, slight as it is, it appears that Forskahl’s name does not refer to
Isolepis uninodis, but may refer to Scirpus litoralis.

From a specimen collected in Ceylon by Koenig, Retzius described a
species which he also called Scirpus lateralis, apparently in ignorance of
Forskahl’s earlier use of the name. Gmelin altered Retzius’ name to Scirpus
lateriflorus, apparently because it was a later homonym. Scirpus lateralis
Retz. was referred to Isolepis supina (L.) R.Br. (Scirpus supinus L.) by
Nees (1834) and to Scirpus supinus L. by C. E. C. Fischer (1932), but other-
wise it and S. lateriflorus appear to have been usually ignored.

Scirpus erectus Poir. was based on a specimen collected by du Petit
Thouars in Mauritius (Madagascar was quoted in the original description,
but the locality on the label is ££ He de France ”). This name was misapplied
by C. B. Clarke (1894 and elsewhere) and by Beetle (1942) to a species
quite distinct from that represented by Poiret’s type. Neither Clarke nor
Beetle included Mauritius in the range of S. erectus, yet Clarke saw Poiret’s
type and annotated it ££ Scirpus supinus Linn. var. y uninodis (sp.) Delile.”

86 PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

Isolepis uninodis Delile was based on an Egyptian plant. C. B. Clarke
(1894, 1902) treated it as a variety of Scirpus supinus and the ternary
combination has been commonly cited as S. supinus L. var. uninodis (Delile)
C. B. Clarke in Hook.f. FI. Brit. Ind. 6 : 656 (1894), although the com-
bination had been published earlier, though rather carelessly, by Ascherson
and Schweinfurth (1887). Chermezon (1931) treated Scirpus erectus and
Isolepis uninodis as synonymous and pointed out that Clarke had misapplied
Poiret’s name to a species of which the correct name is S. juncoides Roxb.
In the synonymy, Chermezon quoted “ Trabut ” as the author of the
combination Scirpus uninodis , but no evidence for this has been found by
me or by Mr. E. Nelmes who kindly made a search for it. Beetle treated
Isolepis uninodis as a distinct species of Scirpus, citing the name as Scirpus
uninodis (Delile) Boiss. FI. Orient. 5 : 380 (1884), but in actual fact Boissier
treated it as a variety of S. supinus (S. supinus L. var. digynus Boiss.)
and cited as a synonym (the basonym) “ S. uninodis Del. Descr. Eg. p. 132
sub Isolepide tab. 6, fig. 1.” Actually, Beetle himself made the transfer
and he correctly cited the author in a later paper (Beetle, 1945) but without
reference to the place of publication. He gave “ Eurasia ” as the range of the
species, although Egypt is the type locality, and followed C. B. Clarke in
misapplying the name S. erectus to S. juncoides. I followed Beetle by
citing S. uninodis (Delile) Boiss. when S. dissachanthus was described and
its affinities discussed (Blake 1946).

“ Isolepis ambigua Steud. ! in Zoll. Verz. Ind. Archip. heft 2, 62 ” was
cited by C. B. Clarke (1907) as a synonym of S. supinus var. uninodis ; this
is a nomen nudum, while the description of I. ambigua Steud. Syn. Cyp. 91
refers to a very different species from those discussed in this paper.

Isolepis oryzetorum Steud. was based on specimens from Java ; the
epithet was spelled “ oryectorum ,” but this is an evident “ unintentional
orthographic error ” (there are other misprints on the page) and the epithet
was correctly spelled in the index.

Among the numerous specimens and other material examined, the
following were of particular importance in interpreting the names mentioned
above: —

The type of Scirpus lateralis Retz. and consequently of S. lateriflorus
Gmel. (ld) ; a rubbing and piece of the type of Scirpus erectus Poir. (p) ; a
tracing of Defile’s excellent plate and a copy of his elaborate description
of Isolepis uninodis (bri) on which name Scirpus supinus var. digynus
must be typified ; topotypes of Isolepis oryzetorum, Scirpus luzonensis,
Scirpus supinus and Scirpus timorensis ; and the original figure and descrip-
tion of Scirpus erectogracilis. Three distinctive species can be [recognised,
all with slender, non- septate culms and very long lower involucral bracts.
They may be distinguished as follows : —

Calms nodeless above the base, without flowers in the leaf-sheaths ;
involucral bract solitary ; spikelets always sessile, usually few,
rarely solitary ; glumes many-striate with glabrous margins ; nut
ob ovate in outline :

Annual plants with black, rather sharply trigonous, prominently
transversely w;avy-ridged nuts and no hypogynous
bristles ; leaf-blades well-developed . . . . . . . . S. supinus

Perennial plants with brown, plano-convex, faintly wavy-
ridged nut and 6 well-developed, retrorsely scabrous,
hypogynous bristles ; leaf-blades rudimentary . . . . S. juncoides

CYPERACEAE AND GRAMINEAE.

Culms with a node well above the base ; leaf-blades rudimentary ;
solitary flowers in the leaf-sheaths with long-exserted stigmas ;

2 involucral bracts usually present, the second one short ; some
spikelets peduncled but the peduncles sometimes very short ;
glumes 3-1-nerved, prominently keeled, minutely ciliolate on
the upper margins ; nut suborb icular in outline, plano-convex to
trigonous, black when ripe, faintly to conspicuously transversely
wavy-ridged . . . . . . . . . . . . . . . . S. lateriflorus

S. supinus appears to be confined to Europe ; its complete synonymy
has not yet been worked out. The synonymy and distribution of the other
two species follow : —

Scirpus lateriflorus Gmel. Syst. 127 (1791).

Scirpus lateralis Retz. Obs. 4 : 12 (1786), non Forsk. (1775).

Scirpus erectus Poir. Encycl. 6 : 761 (1804).

Scirpus polycoleus Notaris in Ann. Sc. Nat. Ill, 9 : 326 (1848).

Scirpus supinus L. var. digynus Boiss. FI. Orient. 5 : 380 (1884), non
Boeck. (1870).

Scirpus supinus L. var. uninodis (Delile) Asch. & Schweinf. 111.
FI. d’%ypte 157 (1887).

Scirpus supinus L. var. uninodis (Delile) C. B. Clarke in Hook.f. FI.
Brit. Ind. 6 : 656 (1894).

Scirpus erectogracilis Hayata Ic. PI. Formos. 6 : 114 (1916).

Scirpus oryzetorum (Steud.) Ohwi in Mem. Coll. Sci. Kyoto Imp.
Univ. ser. B, 18 (1) : ? (1944).

Scirpus uninodis (Delile) Beetle in Amer. J. Bot. 29 : 656 (1942)
( author wrongly cited ) et in Amer. Midi. Nat. 34 : 734 (1945).

Isolepis uninodis Delile FI. Egypt. 8, t. 6, jig. 1 (1812).

Isolepis polycolea (Notaris) Steud. Syn. Cyp. 95 (1855).

Isolepis oryzetorum Steud. Syn. Cyp. 96 (1855) (“ oryectorum ”).

? Isolepis proxima Steud. Syn. Cyp. 95 (1855).

S. lateriflorus is widely spread in the tropics and subtropics of Africa,
Asia and Australia, chiefly in rice-fields, at the edges of swamps and streams
and on the beds of drying-out lagoons and water- courses. The following-
specimens have been examined in various herbaria : —

AFRICA. — Egypt (tracing of Defile’s figure of Isolepis uninodis
Defile). Mauritius, du Petit Thouars (type of Scirpus erectus Poir.).

ASIA. — India : Eastern India, Wight 2893. Ceylon : Ganorawa, in
rice-field, Sept. 1926, Alston 318 ; without definite locality, Koenig (type
of Scirpus lateralis Retz. and S. lateriflorus Gmel.). Formosa : (Original
figure of S. erectogracilis Hayata). Malaya : Kedah, Kepala Batas, Nov.
1941, Corner in Singapore Field No. 38108; Langkaw, rice-fields near
Kuah, Nov. 1941, Corner in Singapore Field No. 37975. Philippine Islands:
Pangasinan Province : Alaminos, Dec. 1922, McGregor in Bureau of Science
No. 41456. Bulacan Province : Manila and vicinity, Oct. 1913, Ramos in
Bureau of Science No. 21970, and without exact locality, Sept. 1913, Ramos
in Bureau of Science No. 1441 partly. Java : Without definite locality,
Ploem ; Geneng, 28th May 1913, Haarsveld ; Redzoso, E. of Pasoeroean,
10th June 1924, Kooper ; (topotypes of Isolepis oryzetorum Steud.).
Soedhono : Without definite locality and without collector’s name, 19th
June 1925. Lombok : Pangantap, coast, July 1909, Griindler 2377.

88 PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

AUSTRALIA. — Western Australia : Kimberley Division : Isdell

R. , 5 miles below Mt. Bartlett, July 1905, Fitzgerald ; Lennard R., 10

miles above junction of Barker R., May 1905, Fitzgerald 588. Northern
Territory : Fitzmaurice R., Oct. 1855, Mueller ; 12° 43' S., 131° 30' E.,
dried-out depressions at edge of lagoon, 17th Sept. 1946, Blake 17033 ; near
Rum Jungle and Finniss R., about 12° 59' S., 130° 58' E., wet sandy creek
bank, 55 m., 8th August 1946, Blake 16728 ; Mary R., about 13° 5' S. and
131° 47' E., muddy bank, about 55 m., 28th Sept. 1946, Blake 17088 ; about
14° 7' S., 131° 16'-18' E., edge of open swamp, about 45 m., 1st July 1946r
Blake 16273. Queensland : Burke District : Near Burketown, June
1943, Whitehouse ; near Normanton, edge of fresh- water lagoon, 19th
August 1936, Blake 12626 ; between Norman and Gilbert Rivers, Gulliver .
Cook District : Endeavour R., Banks cb Solander. North Kennedy District :
Near Woodstock, S. of Townsville, on mud in drying-out Melaleuca swamp,
23rd Sept. 1950, Blake 18510 ; Pentland, edge of tank, 390 m., 12th June
1934, Blake 6148. Mitchell District : Geera, E. of Barcaldine, in fresh
water at 270 m., 29th Nov. 1935, Blake 10337, and in wet places at end of
bore-drain, 270 m., 6th Dec. 1935, Blake 10363. New South Wales : North
Coast : Richmond R., Woolls. Central Coast : Centennial Park, Sydney,.
Feb. 1916, Hamilton. Victoria : Mallee : Lake Lalbert, Mueller

Dimboola, Jan. 1895, Reader.

Of the names cited at the beginning of this discussion, S. hallii Gray,.

S. supinus var. hallii (Gray) Gray and S. uninodis var. hallii (Gray) Beetle
refer to a plant of the eastern United States. Beetle stressed the sharp
ridges on the nut and its prominent mucro ; it is evidently very close to
S. lateriflorus , of which some Australian specimens have sharply ridged
nuts, but I have not seen sufficient American material to form an independent
opinion on its status.

Scirpus juncoides Roxb. Hort. Bengal. 81 (1814), nomen nudum ; FI. Ind.

1 : 228 (1820), ed. Carey 1 : 216 (1832) ; non Willd. ex Kunth

(1837).

Scirpus luzonensis Presl Rel. Haenk. 1 : 193 (1828).

Scirpus junciformis Nees in Wight Contrib. Bot. Ind. 112 (1834) ; non
Retz. (1791) nec (H. B. K.) Poir. (1817).

Scirpus timorensis Kunth Enum. 2 : 162 (1837).

Scirpus supinus L. var. elatior Boeck. in Linnaea 36 : 700 (1870).

Eleochar is juncoides (Roxb.) Schult. Mant. 2 : 90 (1824).

Isolepis (?) juncoides (Roxb.) Miq. FI. Ind. Bat. 3 : 312 (1856).

This is the plant for which the name Scirpus erectus Poir. has been
commonly used. It is a stouter plant than the species to which this name
was originally applied (= S. lateriflorus Gmel.), with few, larger, ovoid,
terete, sessile spikelets, tightly appressed, striate glumes keeled only near
the broadly rounded and more or less mucronate tip and glabrous on the
margins, well- developed hypogynous bristles, and more or less plano-convex
brown nut which is only faintly rugose. It is widely spread and apparently
common from India and China to Malaysia as far east as Timor.

Seleria sumatrensis Retz. Obs. 5 : 19 (1789).

Queensland. — Cook District : Yarrabah near Cairns, in swamp forest*
28th June 1935, Blake 9649 ; Innisfail, in Pandanus swamp, about 4*5 m...
28th Nov. 1941, Blake 14467.

CYPERACEAE AND GRAMINEAE.

New for Australia ; widely spread through Malaysia to Ceylon and
India, but not yet known from New Guinea. The type (ld) consists of an
inflorescence with some immature fruit ; the Australian specimens have
paler spikelets. The species belongs to a group of closely allied coarse
species which have the leaves in the middle part of the stem and the lower
bracts clustered in false- whorls of 3, large inflorescences of several partial
panicles, unisexual spikelets, and a 3-lobed disc ; S. sumatrensis is
distinguished by the tall disc which is from one-half to two-thirds as high
as the nut.

Scleria terrestris (L.) Fassett in Rhodora, 26 : 159 (1924).

Queensland : Cook District : Innisfail, in Pandanus swamp, about

4- 5 m., 28th Nov. 1941, Blake 14468.

New for Australia ; widely spread through Malaysia to China and
Ceylon. It is a coarse species with scattered leaves, unisexual spikelets
and a small disc with rounded short lobes.

Scleria poaeformis Retz. Obs. 4 : 13 (1786).

Scleria oryzoides Presl Rel. Haenk. 1 : 201 (1830) ; Benth. FI. Austral.

7 : 432 (1878).

Queensland. — North Kennedy District : Cardwell, in Melaleuca

swamps about sea-level, dominating with Lepironia articulata the wetter
parts, 26th Sept. 1935, Blake 9700 ; Ingham, fairly common in open swamp,
5th Dec. 1942, Blake 14787.

New for Queensland ; previously reported from the Northern Territory
by Bentham, l.c., under the name of S. oryzoides Presl. The type (ld)
consists of an inflorescence only, which is matched by those of the Queensland
specimens. The stout stems are rather distantly produced from a long-
creeping rhizome and bear scattered, broad, more or less obtuse leaves
and solitary, terminal, decompound panicles of chiefly male spikelets ;
female spikelets occur only towards the base of the branches in the lower
part of the panicles. It is widely spread through Malaysia to India.

Gramineae.

Stipa nodosa S. T. Blake ; species nova, afflnis S. scabrae Lindl., sed
culmis plurinodibus, foliis basalibus perpaucis, ligulis brevioribus
glabrisque praecipue differt.

Gramen perenne, caespitosum, circa 1 m. altum. Culmi erecti, graciles,
teretes, duri, leviter striati, dense granuloso-asperuli, glabri, pro more

5- 8-nodes nodis incrassatis, simplices vel parce ramosi, basi cataphyllis
paucis stramineis plus minusve sericeis cincti ; innovationes perpaucae.
Folia plerumque caulina, basalia perpauca ; vaginae arctae, convolutae,
striatae, asperulae, margine exteriore ciliatae, ore barbatae, nodis glabrae,
internodiis breviores ; auriculae incrassatae ; ligulae truncatae, glabrae,
0-5-0-75 mm. longae ; laminae rigidae, setaceae, convolutae vel involutae,
acutissimae, minute scabridae, usque ad 30 cm. longae, explanatae usque
ad 3*5 mm. latae, sursum nervis scabridae, inferiores gradatim minores.
Panicula exserta, sublaxiflora, linearis, 20-30 cm. longa (aristis exclusis) et
4-6 cm. lata ; axis communis pro majore parte teres laevisque, parte
superiore plus minusve angulosa scabra, nodis (praecipue inferioribus)
albo-pilosa ; rami ad nodum quemque plures, unilateraliter fasciculati,
tenuiter filiformes, suberecti vel patentes, fere ad medium nudi, longiores
usque ad 6 cm. longi iterum ramulosi ; pedicelli sursum scabridi, 4-10 mm.

90 PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

longi. Spiculae lineares, purpurascentes. Glumae subaequales, 7-8 mm.
longae, tenuiter membranaceae, lanceolatae, acutissime acuminatae, glabrae,
carina percurrente sursum vix scabridae ceterum laeves ; inferior 3-nervis
nervis lateralibus brevibus ; superior parum (usque ad 1 mm.) brevior,
5-nervis, nervis exterioribus brevioribus, intermediis cum nervo mediano
supra medium anastomosantibus. Lemma demum fuscum, linear i-fusiforme,
callo incluso 5-5-5 mm. longum, 0-7-0-8 mm. latum, puncticulatum, albo-
pilosum, apice minute lobulato nudum ; callus 1-3-2 mm. longus, albo-
barbatus, apice curvatus pungens. Arista gracilis, 45-50 mm. longa,
scabrida ; columna 11-5-13 mm. longa, sursum flexuosa ; seta falcata,
gracillima. Palea lemmate fere aequilonga, carinis longe pilosa. Plate V.

South Australia. — Flinders Ranges : In Parachilna Gorge between
Blinman and Parachilna, at edge of Parachilna Spring, about 225 m., 1st
Sept. 1946, Blake 16914 (type) ; ENE. of Port Augusta in Pichirichi Pass,
in mallee (chiefly Eucalyptus qracilis) - Kochia open scrub, 29th August
1946, Blake 16863.

Stipa nodosa closely resembles S. scabra in the structure of the
inflorescence and spikelet, but it is entirely different in habit. The hard
culms with usually 5-8 nodes and paucity of basal leaves and innovations
in S. nodosa contrast sharply with the 3-4-noded, softer culms of S. scabra
with their abundance of basal leaves and innovations ; the leaf-blades
are less scabrous and more rigid, the ligule is much shorter and quite
glabrous (not hairy on the back), the panicle is exserted rather early and
not permanently included at its base in the uppermost leaf- sheath as in
S. scabra , and the glumes and awns are slightly shorter. Each of the col-
lections cited consists of several sheets with panicles in various stages of
development. The plants in Pichirichi Pass were associated with S. scabra
{Blake 16864).

Stipa brachystephana S. T. Blake ; species nova, affinis S. aristiglumi
F. Muell., S. bigeniculatae Hughes et S. blackii C. E. Hubbard, sed
ab omnibus ligulis elongatis glabris atque aristis brevioribus differt.

Gramen perenne, caespitosum, circa 70-90 cm. altum. Culmi erecti,
teretes, leviter striatuli, dense granuloso-asperuli, glabri, 3-4-nodes,
simplices, basi cataphyllis subsericeis fulvis vel brunnescentibus cincti.
Foliorum vaginae arctae, convolutae, glabrae, laeves vel minute asperulae,
superiores vel omnes internodiis multo breviores ; ligulae rigidae, striatae,
glabrae, apice integrae vel saepe emarginatae vel tandem bifidae, 4-7 mm.
longae, eae foliorum basalium gradatim breviores ; laminae setaceae, arete
convolutae quasi teretes, rigidiusculae, apice acutissimae sed vix pungentes,
suberectae, longissimae, inferiores medium culmum superantes, superiores
paniculam attingentes, minute asperulae vel laeves, supra pilosulae,
marginibus scabridae, explanatae 0-8-1 mm. latae, eae foliorum basalium
gradatim breviores. Panicula angusta, sublaxiflora, tandem longe exserta,
aristis exclusis 13-15 cm. longa at T5-2-5 cm. lata ; axis communis inferne
compressa sublaevis, sursum angulosa scabrida ; rami terni vel bini, usque
ad 3-5 cm. longi, plus minusve erecti, usque ad medium pro more nudi,
sursum parum divisi pauciflori ; pedicelli compresso-filiformes, scabri, 2-8
mm. longi. Spiculae primum fusiformes tandem plus minusve hiantes,
purpurascentes. Glumae firme membranaceae, margines versus hyalinae,
5-nerves nervo medio plus minusve percurrente sursum scabro, 2 lateralibus
circa median! glumam evanescentibus, 2 exterioribus brevibus, sursum
dorso scabridae, dorso admodum incurvae, explanatae lanceolatae,
acuminatae saepe cuspidatae ; inferior 11-12 mm. longa ; superior 9-10 mm.

CYPERACEAE AND GR AMINE AE.

tonga minus acuminata. Lemma oblanceolato-fusiforme, apice subabrupte
acuminatum, apice integrum pilis albis usque ad 0*9 mm. longis coronatum,
ceterum pilis longis albis dense vestitum, callo incluso 6*5-7 mm. longum,
circa 1*2 mm. latum ; callus curvatus pungens, 1-6-1-8 mm. longus, pilis
albis barbatus. Arista bigeniculata, scabra, 2*3-2*6 cm. longa ; seta recta,
columnam fere adaequans vel quam ea distincte brevior. Palea lemma
adaequans. Plate VI.

South Australia. — Flinders Ranges : Port Germein Pass, steep

stony hillsides with scattered eucalypts and Xanthorrhoea sp., about 210 m.,
29th August 1946, Blake 16858.

On account of the turgid spikelets with firm, prominently 5-nerved,
bulging and upwardly incurved glumes, the prominently bearded tip of the
lemma and the bigeniculate awn, this species appears to be most closely
related to S. aristiglumis F. Muell., S . bigeniculata Hughes and S. blackii
;4C. E. Hubbard, but the very long glabrous ligule (sometimes split to the
base) and relatively short awns and panicle-branches sharply distinguish it
from all three. The glabrous nodes further distinguish it from S. bigeniculata
and S. blackii and the nearly smooth and glabrous leaves and shorter tuft
•of hairs at the apex of the lemma still further distinguish it from S. blackii.
.8. br achy Stephana and S. blackii were found growing in association, but the
hairy leaves and longer panicle-branches of 8. blackii provided a ready
means of distinguishing them in the field.

Echinochloa siagnina (Retz.) Beauv. Agrost. 161 (1812).

Panicum stagninum Retz. Obs. 5 : 17 (1789).

Western Australia. — Kimberley Division : Milligan’s Lagoon near
Wyndham, 10th April 1950, Langfield 211.

Queensland. — Cook District : Forest Home Station, very abundant
in swamps, April 1931, Brass 1886.

New for Australia ; widely distributed in the tropics of the Eastern
Hemisphere and considered an excellent pasture grass in Tropical Africa.
The Australian specimens differ from the type (S. India, Koenig — ld) in
having a scarcely hispid lower glume and a much longer awn to the lower
lemma, but both these characters are variable in this species.

Echinochloa walteri (Pursh) Heller Cat. N. Amer. PI. ed. 2 : 21 (1900).

Panicum hirtellum Walt. FI. Carol. 72 (1788), non L. (1759).

Panicum walteri Pursh FI. Amer. Sept. 66 (1814).

Queensland. — Wide Bay District : Stony Creek near Bundaberg, 2nd
Jan. 1939, Goy Smith 631 ; Bingera Weir near Bundaberg, along river
bank, 30th Dec. 1937, Smith 407 ; near Mapleton, on more or less open
muddy bank of small stream about 150 m., 14th April 1940, Blake 14176.
Moreton District : North Pine R., Petrie, in mud at water’s edge, 26th
Dec. 1930, Blake 91 : Brisbane R., Bailey ; Holland Park, near Brisbane,
common on outskirts of shallow waterhole full of Leersia hexandra and
Phragmites communis, 15th Feb. 1938, Smith 290 ; Ekibin, Brisbane, 8th
May 1909, White ; Samford Creek, common in soft mud on edge of creek,
29th March 1936, Goy 115 ; Enoggera, 17th May 1915, White ; Wellington
Point, March 1916, White ; Currumbin, in a heap of road sand by the
roadside, 10th Dec. 1932, White 8739 ; foot of Mt. Gipps (McPherson
Range), in a gully, about 210 m., 11th April 1941, Blake 14319. Darling
Downs District : “ Merivale ” near Injune (no collector’s name).

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

New South Wales. — North Coast : Lynch’s Creek, in wet gullies in
Eucalyptus forest, 15th March 1944, White 12599.

New for Australia ; native of the eastern United States. The
Australian specimens correspond to forma laevigata Wiegand with almost
glabrous sheaths.

The Australian species of Echinochloa may be distinguished as.
follows : —

Ligule represented by a fringe of hairs :

Spikelets much narrower than long, rather gradually
acuminate and distinctly awned

Spikelets nearly as broad as long, abruptly acuminate
and cuspidate but not awned

Ligule represented by a discoloured zone only :

Lower glume and upper lemma equally acute or
cuspidulate ; racemes 3-4-seriate, ± erect

Lower glume and upper lemma cuspidate or produced
into an awn, the latter more so than the former ;
racemes d: spreading :

Inflorescence d= erect, with awns up to 10 mm.
long or spikelets awnless ; leaf-sheaths with a
few long hairs on the margins or glabrous :

Spikelets ± awned, ± 2-seriate ; racemes
usually straight

Spikelets awnless, 4-seriate ; racemes up-
curved .. .. E.crus-galli var. frumentaceaW. F. Wight

Inflorescence dz nodding, the spikelets partly
hidden by awns 10-25 mm. long ; leaf-sheaths

puberulous on the collar . . . . . . . . E. walteri (Pursh) Heller

Panicum fulgidum Hughes in Kew Bull. 1923 : 323 (1923).

Panicum bicolor R.Br. Prodr. 191 (1810), non Moench. (1794).

Panieum bicoloratum S. T. Blake in Proc. Roy. Soc. Queensl. 59 : 158
(1948).

Panicum bicoloratum was proposed as a new name because the epithet
chosen by Hughes when renaming P. bicolor R.Br. “ had already been
used by Stapf for another species of the genus.” This, however, is incorrect ;
the name published by Stapf was Panicum fulgens, not Panicum fulgidum ,,
and Panicum bicoloratum is therefore a superfluous name. Mr. C. E.
Hubbard, of the Kew Herbarium, kindly drew my attention to the error.

Paspalidium Stapf in Prain FI. Trop. Afr. 9 : 15 (1917), in clavi, et 582
(1920), descr.

The name Paspalidium was proposed for a group of species previously
included in Panicum and still so treated by some American botanists. The
genus has a fairly distinctive facies, and it resembles Setaria rather than
Panicum in the structure of the spikelet and the sterile tip of the branches
of the inflorescence. The genus is widely distributed over the warmer
parts of the world and is particularly well- developed in Australia where at
least 21 of the 30 or so known species have been found. An account of the
Australian species is in preparation, but some new records and descriptions
of new species are given here ; for the latter, only the types and general
range are cited.

E. stagnina (Retz.) Beauv.
E. turneriana Domin

E. colonum (L.) Link

E. crus-galh (L.) Beauv..

CYPERACEAE AND GRAMINEAE.

Paspalidium flavidum (Retz.) A. Camus in Lecomte FL Gen. Indo-Chine
7 : 419(1922).

Panicum flavidum Retz. Obs. 4 : 15 (1786).

Queensland. — Cook District : Mareeba, open sandy ground, 375 m.,
18th June 1935, Blake 9469. North Kennedy District : Mt. Julian, Michael
1405. Port Curtis District: Rosedale, uncommon, Dovey G.30; Rosedale, only
on creek flats, 21st Jan. 1934, Dovey 459. Wide Bay District : Fraser Island,
between Ungowa and the Forestry Station, in mixed forest on sand, 26th
August 1941, Blake 14397 ; Fraser Island, Oct. 1921, White ; Noosa, on sandy
track through open forest above the beach, 15th July 1943, D. A. & L. S.
Smith. Moreton District : Coolum, hillsides in open forest on sand, 15th April
1938, Blake 13747 ; Maroochydore, April 1916, White ; Bribie Island, 10th
April 1938, White ; Stradbroke Island, March 1916, Bick & White ; Enoggera
Range, 1st May 1916, White ; Sunnybank, Dec. 1916, White ; Sunnybank,
open forest on loose sandy soil, 7th Dec. 1933, Blake 4997 ; Sunnybank, 15th
Feb. 1938, L. S. Smith 347 ; Brisbane, shady corner in University grounds,
28th April 1932, Blake 266 ; Canungra, on southern slopes of Mt. Tamborine,
in Eucalyptus forest, 240 m. and upwards, 26th March 1937, Blake 12874.

New South Wales. — North Coast : Between Kingscliff and Norries
Head, among Imperata and Pteridium in mixed open forest on slopes of low
sandridges behind the coast, 19th April 1950, Blake 18495.

New for Australia, as all previous records from here appear to be based
on misidentifications. It was definitely recognised by C. E. Hubbard when
he sorted the material in the Queensland Herbarium in 1930-1. The
Australian material agrees well with the type collected in Ceylon by Koenig
(ld), but has somewhat longer leaves ; the length of the spikelets varies
from 2-6 to 3 mm. (2*6-2*9 mm. on the type), which is somewhat greater
than the length given in most published descriptions of the species. Perhaps
other species have been confused with it in Ceylon and India as in Australia.
The following names and references must be considered in explaining the
confusion with other Australian species : —

Panicum hrizoides L. ; F. Muell. Fragm. 8 : 189 (1874).

Panicum distans Trin. Spec. Gram. t. 172 (1829) ; non Willd. ex Spreng.
Syst. Veg. 1 : 305 (1805), pro syn.

Panicum flavidum Retz. Obs. 4 : 15 (1786) ; R.Br. Prodr. 190 (1810) ;
Benth. FI. Austral. 7 : 474 (1878) ; Domin in Biblioth. Bot. 85 : 300
(1915).

Panicum flavidum Retz. var. jubiflorum (Trin.) Domin in Biblioth. Bot.
85 : 300 (1915).

Panicum flavidum Retz. var. orarium Domin in Biblioth. Bot. 85 : 300
(1915).

Panicum flavidum Retz. var. tenuius Benth. FI. Austral. 7 : 474 (1878)
(“ tenuior ”) ; Domin in Biblioth. Bot. 85 : 300 (1915).

Panicum globoideum Domin in Fedde Repert. Nov. Sp. 10 : 119 (1911).

Panicum gracile R.Br. Prodr. 190 (1810) ; Benth. FI. Austral. 7 : 475
(1878).

Panicum jubiflorum Trin. Gram. Panic. Dissert. 2 : 150 (1826).

94 PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

Panicum paractaenum Kunth Enum. 1 : 134 (1833).

Panicum retiglume Domin in Fedde Repert. Nov. Sp. 10: 119 (1911).

Paractaenum novae-hollandiae Beauv. Agrost. 47, t. 10,/. 6 (1812) ; Hughes
in Kew Bull. 1923 : 287-9 (1923).

Paspalidum distans (Trin.) Hughes in Kew Bull. 1923 : 317 (1923).

Paspalidium flavidum (Retz.) A. Camus in Lecomte FI. Gen. Indo-Chine
1: 119 (1922).

Paspalidium globoideum (Domin) Hughes in Kew Bull. 1923 : 317 (1923).
Paspalidium gracile (R.Br.) Hughes in Kew Bull. 1923 : 318 (1923).
Paspalidium jubiflorum (Trin.) Hughes in Kew Bull. 1923 : 317 (1923).
Paspalidium retiglume (Domin) Hughes in Kew Bull. 1923 : 317 (1923).

The first reference to the occurrence of Paspalidium flavidum in Aus-
tralia was made by R. Brown ( Panicum flavidum ) ; he indicated that he
saw living plants on the tropical coast, but I have seen no further reference
to these specimens, unless they are the ones referred to Paspalidium
jubiflorum by Hughes. F. Mueller treated Panicum flavidum, P. distans,
P. jubiflorum and P. paractaenum as synonyms of P. brizoides L. which
appears to be truly a synonym of Echinochloa colonum (L.) Link, but which
has been misapplied to P. flavidum. Bentham cited four Australian col-
lections under Panicum flavidum ; of these : one (Upper Victoria R., Mueller )
became the type of Panicum retiglume, two (Peak Downs, Burkitt, and
Springsure, Wuth) became syntypes of P. globoideum, while the fourth
also appears to belong to P. globoideum (to judge from later references).
Panicum flavidum var. tenuius was based on several specimens, some of
which were later distributed by Hughes between Paspalidium jubiflorum
and Paspalidium distans. Bentham also treated Panicum jubiflorum, P.
distans and Paractaenum novae-hollandiae as synonyms of P. gracile. Domin
(1915) referred only one collection to P. flavidum, his own from Winton,
but from geographical considerations it must belong to P. jubiflorum.
Under Panicum flavidum van. jubiflorum he synonymised Panicum jubiflorum
and P. distans, but the only specimen cited came from a wet coastal locality
and cannot be P. jubiflorum which belongs to the drier inland regions.
From the notes given, P. flavidum var. orarium appears to be P. distans.

Paractaenum novae-hollandiae ( Panicum paractaenum ) is generically
distinct from Paspalidium (Hughes, 1923). The species confused with
Paspalidium flavidum may be distinguished by the following key: —

Rhaehis of racemes articulate with the main axis, the

racemes falling entire . . . . . . . . . . Paractaenum novae-liollandiae

Rhaehis of racemes not articulated with the main axis,
long persistent ; spikelets falling separately and singly
{ Paspalidium ) :

Upper glume 9-13-nerved with the nerves anasto-
mosing by means of cross-veins, at least in the upper
part :

Spikelets 2-2*7 mm. wide ; nerves anasto-
mosing only in the upper part ; densely tufted

perennial . . . . . . . . . . P. globoideum

Spikelets 1*4- 1*5 mm. wide ; nerves anasto-
mosing almost throughout ; annual .... P. retiglume

CYPERACEAE AND GRAMINEAE.

Upper glume 5-7 (rarely 9) -nerved ; nerves free or
uniting at or very close to the tip :

All or nearly all the racemes with the spike] ets
closely and evenly biseriate on pedicels
02-05 mm. long, those of each series more or
less contiguous ; racemes nearly always
simple :

Spikelets about twice as long as wTide ; all
or nearly all the racemes longer than the
internodes of the main axis ; leaf- sheaths
glabrous throughout

Spikelets about 1*5- 1-7 times as long as
wide ; lower racemes much shorter than
the internodes of the main axis ; leaf-
sheaths more or less ciliate on at least one
margin, particularly near the top :

Spikelets 2-6-3 mm. long, 1-4-1-6 mm.
wide, widely spreading from the
rhachis, those of each series con-
tiguous for about half their length,
not strongly curved in profile

Spikelets 2-2-5 mm. long, 1-1-1-25
mm. wide, obliquely spreading, those
of each series contiguous only by the
base of the one above and the tip of
the one below

Racemes with the spikelets loosely arranged or
irregularly biseriate, slightly contiguous to
distant on pedicels 0-25-5 mm. long ; at least
the lower racemes with 1 or more 1 -flowered
branches . .

P. jubiflorum

P. flavidum

P. distans

P. gracile

Paspalidium basicla&um Hughes in Kew Bull. 1923 : 318 (1923).

Queensland. — Burke District : Cloncurry, on stony to rocky quartzite
hillsides about 225 m., 7th Nov. 1935, Blake 10123. Gregory North District :
Tranby, 22° 40' S., 142° 25' E., gullies in rugged sandstone ridges, 165-210 m.,
with Triodia sp., 8th May 1936, Blake 11410, 11411 ; Elderslie, W. of
Winton, garden weed, one specimen seen, 27th Oct. 1935, Blake 10006.

South Australia. — Flinders Ranges : Near Blinman, on barren

stony slopes in mixed low scrub, 420-450 m., 31st August 1946, Blake
16904.

New for both Queensland and South Australia.

Paspalidium clementii (Domin) C. E. Hubbard in Kew Bull. 1934 : 447
(1934).

Panicum clementii Domin in J. Linn. Soc. Bot. 41 : 272 (1912).

Queensland. — Gregory North District : Duchess, in rock crevice on
low rugged hill, about 390 m., 18th May 1936, Blake 11531 ; near Boulia,
on low stony ridge with scattered Acacia spp., etc., 24th July 1936, Blake
12379 ; Tranby, 22° 40' S., 142° 25' E., gullies in rugged sandstone ridges,
165-210 m., with Triodia sp., 8th May 1936, Blake 11412 ; Elderslie, W.
of Winton, on stony hillside, 5th June 1936, Blake 11657. Gregory South
District : Betoota, on barren stony desert hills, 17th July 1936, Blake
12177 ; 45 miles W. of Windorah, on scrubby, stony, sandstone ridges,
about 150 m., 14th July 1936, Blake 12125 ; Mt. Howitt Station, about

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

70 miles W. of Eromanga, on barren sandstone hillsides under Acacia
spp., 180-240 m., 3rd July 1936, Blake 11917. North Kennedy District:
Charters Towers, sandstone ridge, Eucalyptus -Acacia forest, 330-360 m.,
11th June 1936, Blake 11705.

New for Queensland.

P. basicladum and P. clementii were hitherto known only from Western
Australia, the former from the south, the latter from the north. The
specimens cited constitute a wide extension of range for both species, and
it is interesting to note that the ranges of the two overlap in western
Queensland, both having been collected together at Tranby and in different
years at Elderslie. They are closely allied annual species which may be
distinguished as follows : —

Spikelets 2-75-3- 1 mm. long, elliptic-oblong, i acute ; upper
floret distinctly attenuated towards the base, coarsely though
shallowly rugose ; culms ± scabrous ; lower racemes mostly
overlapping . . . . . . . . . . . . . . P. basicladum

Spikelets 2-2-4 mm. long, oblong to somewhat elliptic, i obtuse ;
upper floret scarcely attentuated towards the base, faintly and
finely rugose ; culms smooth ; lower racemes ± distant . . P. clementii

Two other annual species are known. P. tabulatum (Hack.) C. E.
Hubbard is distinguished by its filiform leaves and P. rarum (R.Br.) Hughes
by all the racemes reduced to 2 or (more usually) 1 spikelet.

Paspalidium albovillosum S. T. Blake ; species nova affinis P. distanti
(Trin.) Hughes et P. caespitoso C. E. Hubbard ; ab hoc foliis latioribus
ab illo spiculis haud incurvis, ab utroque spiculis paullo minoribus
latioribusque atque foliis et inflorescentiae axi ramisque pilis longis
mollibus vestitis differt.

Gramen perenne, pallide virens vel flavo virens. Culmi caespitosi,
recti, usque ad 40 cm. longi, graciles, molles, laeves, 3-5-nodes, ramosi,
prope inflorescentiain saepe laxe pilosi ceterum glabri. Folia pilis longis
gracilibus mollibus albis e tuberculis ortis dense hirsuta ; vaginae plus
minusve carinatae, arete convolutae vel hiantes, tenues, conspicue nervosae,
margine altero longe ciliatae, dorso sursum hirsutae, nodis glabrae,
inferiores internodiis longiores, superior es breviores ; ligulae ad marginem
augustissimum breviter ciliatum redactae, in toto circa 0*5 mm. longae ;
laminae molles, anguste lineares, breviter acutae, planae vel plus minusve
incurvae, explanatae 1-4 mm. latae, usque ad 15 cm. longae, utrinque
molliter hirsutae, sursum marginibus scabridae. Inflorescentia longe
exserta, 8-15 cm. longa ; axis communis plus minusve triquetra, pro
majore parte vel omnino longe laxeque pilosa (pilis tenuibus mollibus e
tuberculis ortis), sursum angulis minute scabrida. Racemi 6-10,
inferiores plerumque 1-2 cm. longi, internodiis axeos communis breviores,
interdum prope basim ramosi, superiores gradatim breviores internodiis
paullo longiores, simplices ; rhachis flexuosa, circa 0*25 mm. lata,
triquetra, marginibus longe ciliata, seta 0-5-2 mm. longa terminata ;
pedicelli 0-2-2-4 mm. longi, pilosi, apice discoidei, inter se 0-25-1-5 mm.
distantes. Spiculae purpurascentes, biseriatae, contiguae, oblique patentes,
ambitu ellipticae vel ovato-ellipticae vel obovato-ellipticae, late acutae,
prope basim admodum attenuatae, facie depressae a latere visae rectae
(haud incurvae), dorso alte convexae, 1-5-1-9 mm. longae, 1-1-2 mm.
latae, glabrae. Glumae membranaceae, tenuiter nervosae ; inferior
dimidiam spiculam aequans vel fere aequans et ejus basim amplectens,

cyperaceae and gramineae.

oblata, late obtusa, 3-nervis nervis lateralibus cum medio prope apicem
conjunctis ; superior tres usque novem partes spiculae attingens, late
xotundata, 5-7 -nervis nervis subparallelis fere percurrentibus. Anthoecium
inferum sterile : lemma spiculam aequans, explanata late rotundata,

5-nervis ; palea usque circa 0-6 mm. longa vel 0. Anthoecium superum
spiculae apicem attingens, fere ellipticum, utrinque acutum, apice
subacuminatum brevissime apiculatum (apiculo fere recto), dorso le viter
reticulatum minime rugulosum.

Typus : Queensland; Maranoa District, 20 miles W. of Mitchell, dense
Gadellia- Acacia forest, greyish gravelly silt loam, 480 m., 31st March
1936, Blake 10947.

The species is represented by 23 collections from Queensland collected
in the Districts of South Kennedy, Wide Bay, Burnett, Leichhardt,
Warrego, Maranoa, Darling Downs and Moretom It is easily recognised
by the small but broad spikelets and the long, slender, soft, more or less
spreading hairs on the leaf-blades, main axis of the inflorescence, rhachis of
the racemes and the pedicels.

Faspalidium spartellum S. T. Blake ; species nova, affinis P. gracili
(B.Br.) Hughes, sed foliorum vaginis (haud carinatis) laminisque
brevibus, racemis omnibus 1-2-spiculatis, pedicellis perbrevibus,
anthoecio supero basim versus rotundato apice recto dorso vix
ruguloso differt.

Gramen perenne, caespites duros eff ormans. Culmi erecti, circa 50 cm.
alti, graciles, duri, rigidi, teretes, striolati, laeves, glabri, 5-8 -nodes, iterum
ramosi ramis ramulisque patulis. Folia brevia ; vaginae primum convolutae
teretes, demum solutae et apertae, haud carinatae, conspicue nervosae,
internodiis multo breviores, glabri laeves que ; ligulae ad marginem
brevissime ciliolatum in toto circa 0-15 mm. altum redact ae ; laminae
patulae vel deflexae, angustissime lineares, acutissimae, rigidae,
involutae, explanatae usque ad 1 mm. latae, nervis utrinque
scabridae ceterum glabrae laevesque, usque ad 6 cm. longae. Inflorescentia
exserta, pauciflora, subracemiformis, 1 *5-3-5 cm. longa ; axis communis
inferne compresso-canaliculata sursum subtriquetra, omnino scabrida.
Racemi 4-6, erecti, ad spiculas 1-2 redacti, inferiores distantes saepe ramosi,
superiores contigui simplices ; rhachis triquetra, scabrida, circa 0*25 mm.
lata, seta subulata 0*5-1 *5 mm. longa terminata ; pedicelli 0-4-0-6 mm. longi,
scabridi, apice discoidei, usque 1*5 mm. distantes. Spiculae brunnescentes,
erecti, ambitu angustius ellipticae acutae, prope basim constrictae, a latere
visaerectae, 2*9-3 mm. longae, 1* 1-1*2 mm. latae, conspicue nervosae. Glumae
membranaceae ; inferior ovata, acuta, duas partes spiculae aequans vel
fere aequans, valide 3-nervis nervis prope apicem convergentibus ; superior
quattuor partes spiculae aequans, specie acuta sed explanata rotundata,
5-7-nervis, nervis 5 prope apicem convergentibus et ibi cum se unitis ceteris
brevibus. Anthoecium inferum sterile ; lemma glumae superiori simile
sed spicula aequilongum, 7-nerve nervis prope apicem convergentibus cum
se conjunctis ; palea 0. Anthoecium superum spicula aequilongum,
oblongo- ellipticum, acutum, minute apiculatum, piano- con vexum, tenuiter
reticulatum, dorso inferiore inconspicue tenuissimeque rugulosum.

Typus : Queensland; North Kennedy District, W. of Pentland, between
Warrigal and Burra, on slopes of Great Dividing Range, in open forest on
shallow sand overlying sandstone, 450-495 m., 19th October 1935, Blake
9939.

98 PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

Known only from the type- collection, this species is very distinct im
appearance by reason of the wiry branched culms, short and often deflexed.
leaves with short spreading sheaths which at length open out to become^
nearly flat, and short, reduced inflorescences of few racemes with only 1-2
strongly nerved spikelets.

Paspalidium criniforme S. T. Blake ; species nova, affinis P. gracili
(R.Br.) Hughes, sed spiculis paullo minoribus pro rata angustioribus,
anthoecio supero brevissime apiculato basi vix attenuato praecipuo
differt.

Gramen perenne viride. Culmi caespitosi, erecti vel geniculati, circa
10-27 cm. alti, setacei, compressi, striolati, glabri, laeves, 4-7-nodes,
iterum ramosi. Folia pilis longiusculis tenuibus e tuberculis parvis ortis
praedita ; vaginae carinatae, crebre nervosae, laeves, sursum laxe pilosae,
ore barbatae, vel omnino glabrescentes, primo convolutae tandem per
ramos solutae, inferiores internodiis longiores, super iores eis breviores ;
ligulae ad seriem ciliorum vix 0-2 mm. longorum redactae ; laminae
plerumque setaceo-involutae, raro planae et usque 1*5 mm. latae, usque 6-5
cm. longae, utrinque pilosae, supra nervis marginibusque scabridulae.
Inflorescentia fere filiformis, pauciflora, 2*5-6 cm. longa vel interdum
brevior ; axis communis canaliculato-triquetra, sursum minute scabrida.
Racemi 4-6, appressi, inferiores usque 1 cm. longi, saepe ramosi, usque
5 spiculas gerentes, super iores breviores, his vel interdum omnes unispiculati ^
rhachis anguloso-filiformis, circa 0*15 mm. lata, scabrida, leviter flexuosa,
seta 2-4 mm. longa terminata ; pedicelli 1-2 mm. longi, scabriduli, apice
discoidei. Spiculae quasi uniseriatae, pallidae, suberectae glabrae, ambitu
elliptico-lanceolatae vel oblongo-lanceolatae, acutae, acuminato-apiculatae,,
piano- convexae haud incurvae, 2-2*2 mm. longae, 0*9-1 mm. latae, nervis
angustis viridibus conspicue notatae. Glumae membranaceae ; inferior
ovata, acuta, spiculae duas partes aequans vel fere aequans, 3- vel sub-5-
nervis ; superior elliptica, late rotundata, spiculae tres usque quinque
partes aequans, 5- vel sub-7 -nervis nervo mediano percurrenti lateralibus
prope apicem incur vis cum se junctis, eis paris tertii brevibus saepe liberis.
Anthoecium inferum sterile : lemma spiculam aequans, specie apiculatum
sed explanatum ellipticum rotundato-obtusum, 5-nerve, utrinsecus nervum
medianum sulcatum ; palea minuta. Anthoecium superum ova turn,,
acutum, minute apiculatum (apiculo incurvo), dorso alte convexo rugulosum.

Typus : Queensland ; Moreton District, Moggill near Brisbane, oh

dry hillside in mixed open forest, 10th March 1934, Blake 5282.

The species is known from five collections from the Moreton
District of Queensland. The dense tufts of very slender culms and very
fine leaves and the very slender inflorescences of few, small spikelets give
the plant a distinctive appearance. The living plants have a faint but
distinct curry-like odour.

Paspalidium udum S. T. Blake ; species nova, affinis P. punctato (Burm.)
A. Camus (speciei asiaticae), P. paludivago (Hitchc. & Chase) Parodi
(speciei americanae) et P. geminato (Forsk.) Stapf (speciei fere-
cosmotropicae), sed spiculis majoribus, gluma superiore manifesto
5-nervi, anthoecio supero manifeste ruguloso ab omnibus differt.

Gramen perenne, viride. Culmi elongati, usque metrales, basi plus
minusve repentes et nodis inferioribus radicantes, sursum adscendentes,.
molles, striati, laeves, glabri, multinodes, ramosi, inferne usque 7 mm.
crassi. Folia plus minusve conferta ; vaginae primo convolutae mox

CYPERACEAE AND GRAMINEAE.

hiantes, parte superiore carinatae, glabrae laevesque, internodiis
plerumque longiores ; ligulae ad marginem longe ciliatum redactae, in
toto circa 1-5 mm. longae ; laminae anguste lineares, acutae, basim versus
le viter angustatae, planae vel saepius convolutae vel involutae, marginibus
atque nervis supra scabridae ceterum glabrae laevesque, quoad magnitudo
maxime variabiles, saepius 4-8 mm. latae et 5-20 cm. longae. Inflorescentia
angustissima, tandem exserta, 14-22 cm. longa ; axis communis interne
compressa canaliculata laevis, sursum triquetra admodum scabrida.
Racemi 10-16, appressi, inferiores distantes, superiores vel plures imbricati,
omnes simplices, plerumque 1-5-3 cm. longi, supremi plerumque breviores ;
rhachis leviter anfractuosa, triquetra, scabro-ciliata, circa 0-6-0-7 mm. lata,
seta subulata 1-4 mm. longa terminata ; pedicelli cuneati, scabri, 0-2-0-3 mm.
longi, apice plus minusve discoidei, circa 1-1*5 mm. distantes. Spiculae con-
fertae, eleganter biseriatae, fere erectae, eae seriei cuj usque longe imbricatae,
manifestius nervosae, ambitu oblongo-ovatae vel oblongo-ellipticae, acutae,
a latere visae rectae, facie leviter concavae, 3-3-3 mm. longae, 1-2-1-5 mm.
latae. Glumae membranaceae ; inferior quartam vel tertiam partem
spiculae subaequans, oblata, truncata vel admodum rotundata, enervis ;
superior tres quintas vel tres quartas partes spiculae aequans, elliptica,
rotundo-obtusa, conspicue 5 -nervis nervo mediano percurrente ceteris
subparallelis prope marginem incurvis conjunctisque vel nervo transverso
irregulari inconspicuove unitis. Anthoecium inferum sterile : lemma

membranaceum, spicula aequilongum, explanatum oblongo-ellipticum
rotundo-obtusum, 5-nerve nervis sursum convergentibus et juxta marginem
unitis mediano percurrenti ; palea lemmate aequilonga, hyalino-
membranacea, 2-carinata, marginibus late inflexa. Anthoecium superum
spicula aequilongum vel fere aequilongum, ambitu ovatum acute
acuminatum, apiculatum, a latere visum rectum, depresse piano- convexum,
reticulatum et rugulosum ; lemma tenuiter 5-nerve. Antherae 1-2 mm.
longum.

Typus : Northern Territory ; Near Alligator Point, Daly R., 13°

26'-29' S., 130° 26'-27' E., common in damp places with Phyla nodiflora,
15 m., 28th July 1946, Blake 16659.

Known only from the type- collection, this species differs in habit
from other Australian species in the coarse, spongy, creeping and ascending
culms in which it resembles P. geminatum (Forsk.) Stapf (a nearly
cosmotropical species which is the type of the genus), P. punctatum (Burm.)
A. Camus (from Asia) and P. paludivagum (Hitchc. & Chase) Parodi (from
America). The first-mentioned differs from the others in its much smaller,
rather broadly ovate spikelets with an almost quite smooth fertile floret ;
it was based on Panicum geminatum Forsk., the type of which is a specimen
collected at Damietta, Egypt, by Forskahl (c). P. punctatum,
P. paludivagum and P. udum are much alike in general appearance and
characters of the spikelet, but there are minor differences in the size and
shape of the spikelet, the relative length of the upper glume, the number
and distinctness of the nerves of the upper glume and lower lemma, and
in the surface of the fertile floret. P. udum has the largest spikelets with
the most prominent nerves and distinctly rugulose fertile florets ; its upper
glume is 5-nerved and at least usually 3-nerved in the others.

The combination Paspalidium paludivagum has been made
independently on three different occasions : Parodi Gram. Bonar. ed. 3,
85, 89 (1939) ; Henrard in Blumea 3 (3) : 434 (1940) ; and Pilger in Engl.
& Prantl Pflanzenfam. ed. 2, 14e : 29 (1940).

100

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND,

EXPLANATION OF PLATES.

Plate Y.

Stipa nodosa S. T. Blake. Fig. 1, portion of plant, natural size; 2, ligule,
with part of sheath and blade, flattened out X 6 ; 3, transverse section of leaf,
X 30; 4 spikelet, X 3; 5, lower glume, flattened out, X 6; 6, upper glume,
flattened out, X 6 ; 7, floret, greater part of awn removed, X 6 ; 8, part of lemma
with base of awn, flattened out, X 6 ; 9, palea, flattened out, X 6. — From type.

Plate YI.

Stipa brachystephana S. T. Blake. Fig. 1, portion of plant, natural size;
2, ligule with part of sheath and blade flattened out, X 6; 3, transverse section
of leaf X 30; 4, spikelet, X 3; 5, lower glume, flattened out, X 6; 6, upper

glume flattened out, X 6; 7, floret, greater part of awn removed, X 6; 8, tip of
lemma with base of awn, flattened out, X 6; 9, palea, flattened out, X 6. — From
type.

Proc. Koy. Soc. Q ’land, Vol. LXIL, No. 10. Plate

Stipa nodosa ,S. T. Blake.

S.T. BLAKE

Plate VI.

Proc. Roy. Soc. Q’land, Yol. LXII., No. 10.

Stipa hrachy Stephana S. T. Blake.

ST BLAKE

Vol. LXIL, No. 11.

101

NOTES ON SOME AUSTRALIAN COMPOSITAE.

By J. H. Willis, National Herbarium of Victoria, Melbourne.

(With Plate VII and one Text-figure.)

(. Issued separately , 22nd August, 1952.)

Summary.

Two new species of Helichrysum and one of Chthonocephalus are described
from Queensland, New South Wales, and Western Australia respectively, and a
necessary change made in the name of a species of Erechthites , now regarded as
a Senecio. The distributions of four species of Helichrysum are mapped.

Helichrysum eriocephalum J. H. Willis; species nova ex affinitate
H. cordati DC., H. ramosi DC. et H. obovati DC., a quibus differt
foliis supra constanter arachnoideis, inflorescentiis multo minoribus
(in specimine unico), involucri squamis longius lanigeris laminis
albis latioribus plicatisque.

Suffrutex laxus, saltern 50 cm. altus, ramis gracilibus albido-
tomentosis. Folia lanceolata brevissime petiolata, inferiora circiter
5X1 cm., summa minora, distantia, omnia utrinque arachnoideo-
lanuginosa, marginibus revolutis, subtus nervo medio lateralibusque
paucis prominentibus. Inflorescentia densissime lanuginosa terminals
restricta, corymbis confertis in panicula parva laxa dispositis. Capitula
albida subsessilia subhemisphaerica, circa 5 mm. lata, usque 40-flora.
Involucri squamae 30-40, circa 5-seriatae, spathulatae, superiores 2-3
mm. longae, stipite a lana occulto, solum lamina alba brevi (usque ad
1 mm. longa) lata obtusa subplicataque aperta; ob vestem arachnoideo-
lanuginosam intricatam difficillime segregandae. Flores circa 3 mm.
longi, corollae lobis 5 brevibus papulas sparsas ferentes. Pappi setae
circa 20, circa 2-5 mm. longae, scabridae, corollam vix excedentes,
ad apicem incrassatae, ad basin in corona (latiore quam ovario) connatae.
Achaenia immatura breviter cylindrica glabra. Styli ramuli 0-5-1 *0 mm.
longi, ad apices brevissime denseque penicillati. Antherae circa 1-5 mm.
longae, caudis brevibus paulum lobatis; filamentis gracilibus. Pollinis
granula circa 18 mic. diam., regulariter echinulata. — Plate VII, figs.
1-7.

Queensland. — South Kennedy District: Lake Elphinstone, about 100 miles
WSW of Mackay, 1870, Amalie Dietrich 1722 (holotype in hel; clastotype in

BRI.).

Notwithstanding the undesirable practice of describing new species
from single herbarium specimens, I do not hesitate to publish a diagnosis
of this hitherto undescribed plant which differs in so many ways from
other species of Helichrysum. P. Mueller had long ago labelled the
collection “H. Beckleri P.v.M. (var.)”; but it has conspicuous white
and crinkled spreading tips to the involucral bracts, both surfaces of
the rather large lax remote leaves persistently arachnoid-woolly, and
bears very little resemblance to this shrub. In January 1947 the late
Mr. C. T. White, to whom I submitted this puzzling Queensland plant
for examination, reported that he had seen nothing like it and suggested
my describing it as new.

With, apparently, a weak and straggling habit, its white-woolly
branches, moderately large, broad leaves and exceedingly woolly heads
(the bracts difficult to disentangle), H. eriocephalum is obviously allied

102

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

to H. cor datum, H. ramosum and H. obovatum — all Candollean species
in the Section Ozothamnus. II. obovatum of north-eastern New South
Wales differs in having glabrescent leaves on slender petioles and very
minute, non-spreading tips to the involucral bracts. The other two
species are Western Australian and differ in their large, diffuse, leafless
panicles, the bracts much less hairy with narrower white tips (never
crinkled as in the new species), and the more sharply scabrid pappus
bristles which are not fused at the base to form a broad crown. It
would be most interesting to ascertain whether the plant still survives
in the neighbourhood of Lake Blphinstone and what is the nature of
its habitat,

Helichrysum tuckeri F. Muell. ex J. H. Willis; species nova, de forma
foliorum solummodo H. diotophyllo F. Muell. comparabilis, cujus
autem folia basi amplius auriculata, sursum angustiora et
marginibus paulo distantibus subtus tomentosa, capitula ampliora
flores usque 40 tenentia, involucrum nitenti-flavescens, pappi setae
minus incrassatae et achaenia fere sericata sunt; H. diosmifolium
(Vent.) Sweet, quod forma et colore capitulorum speciei novae
simillimum, foliis multo longioribus anguste linearibus
patentioribus, floribus in capitulo pluribus differt.

Frutex rigidus, 60-120 cm. altus, ramis numerosis virgatis gracilibus
tenuissime tomentellis. Folia glabra angusto-oblonga, obtusa sessilia
perbrevia (eireiter 3-4 X 1 mm.), ad basin latiora et brevissime biloba,
erecta appressa conferta, subtus sulco angustissimo percursa, marginibus
replicatis paginam inferam plane occultantibus. Inflorescentia
terminalis, paniculata, fere globularis, 1-3 cm. lata, ex corymbis
capitulorum constructa. Capitula albida, baud nitentia, breviter
peduneulata, primum pilularia demum campanulata, 3-4 mm. lata,
7-12-flora. Involucri squamae 30-40, circa 5-seriatae, integrae, glabrae
vel ad stipitem parcissime lanuginosae; superiores obovato-oblongae,
circa 3 mm. longae, sursum albae, inferne secus stipitem hyalinae ;
inferiores minor es ad formam orbicularern vergentes. Flores circa
2*5 mm. longi, sub anthesi involucrum parum superantes, corollae lobis
5 mmutissime papulatis. Pappi setae corollam aequantes baud
numerosae, 1*5-2 mm. longae, minute scabridae, apicem versus leniter
incrassatae, ad basin fere liberae. Achaenia circa 0*5 mm. longa,
breviter cylindrica, dense papillose scabriuscula. Styli ramuli circa
0*6 mm. longi, ad apices brevissime denseque penicillati. Antherae circa
1 mm. longae, caudis brevibus acuminatis ; filamentis gracilibus.
Pollinis granula circa 18 mic. diam., sparsim echinulata. — Plate VII,
figs. 14-20.

New South Wales. — Western Plains: Lake Cargelligo, Nov., 1915, J. W. Dwyer ;
‘-‘Lachlan Elver/’ 1879, Gerard Tucker (holotype in mel, clastotype in nsw) ;
Shuttleton Dee. 1903, W. Bduerlen; Lachlan to Darling River, G. Day ; Nyngan,
Dee. 1899, W. Bduerlen; Girilambone, Nov., 1890, E. Betche; Pilliga-Gwabegar,
Dec., 1932 H. M. B. Bupp. Central Western Slopes: Kamarah via Bar ell an,
17th Get., 1917, W. B. A. Baker ; Ardlethan, Nov., 1917, J. L. Boorman; Weddin,
Dec. 1899, J. H. Maiden; Wyalong, Oct., 1903, J. L. Boormom; Wyalong to
Barmedman, Mar., 1915, J. W. Dwyer ; Temora, Oct., 1915, J. W. Dwyer ; Dudauman,
10 miles NW. of Cootamundra, Nov., 1917, J. L. Boorman. Southern Western
Slopes: Near Wagga, Wagga (? collector).

! Victoria. — “Wimmera,” 1894, W. D. Matthews.

[All collections available in the National Herbaria at Melbourne, Sydney and

Brisbane are cited here.]

NOTES ON SOME AUSTRALIAN COMPOSITAE.

103

Adopting a manuscript epithet given in 1879 by F. Mueller, who
was the first to recognize the specific distinctiveness of H. tuckeri, I
have drawn freely from his detailed notes accompanying the type
specimen in the Melbourne Herbarium. Apparently it had been
Mueller ’s intention to publish this name, honouring the original collector,
and his epithet is now perpetuated as a mark of respect.

This rigid shrub (to four feet high) has hitherto been confused
with H. diotophyllum F. Muell. which has rather similar appressed
foliage and shares the northern part of its range on the Western
Plains of New South Wales. Leaves of the new species are almost
intermediate between those of II. diotophyllum (with large basal
auricles) and the non-auriculate H. adnatum (DC.) Benth. ; but its
white, pilular flower-heads in small, globoid panicles are quite dissimilar,
more closely approaching those of the much larger-leaved
H. diosmifolium (Vent.) Sweet. All four species occur in New South
Wales and are confined to eastern Australia, between the 25th and 37th
parallels of south latitude (as far as existing records show).
H. diosmifolium has by far the longest range, extending from Mt.
Faraday (NE. corner of Warrego District) and Laguna Bay in Queens-
land to Hervey’s Range near Peak Hill and Mt. Dromedary on the
SE. coast of New South Wales; it is predominantly a plant of better-
watered, hilly country, and in the southern area of distribution mingles
with U. adnatum (from Walcha on and east of the Dividing Range to
Suggan Buggan in far eastern Victoria where several other tableland
species reach their southern limit). II. diotophyllum is co-extensive
with H. diosmifolium in the Darling Downs District — the only two
members of this group in Queensland — while in western New South
Wales (as at Shuttleton) it accompanies H. tuckeri which ranges from
Pilliga southward to Wagga Wagga, with a strong development in the
Wyalong-Temora district.

A collection of H. tuckeri in the Melbourne Herbarium labelled
“Wimmera” (from W. H. Matthews at Stawell Technical College,
1894) is of doubtful origin. It was exhibited by Mueller as new for
Victoria* at the meeting of the Victorian Field Naturalists’ Club in
March, 1894, and is the basis of the record of H. diotophyllum in
Victoria by Ewart (1930). No other specimen of the species is known
from this State and it is highly probable that Matthews either collected
his material during a visit to New South Wales or received it through
some correspondent there — certainly Stawell is far beyond the expected
dryland habitat of H. tuckeri, not to mention the still more northerly
H. diotophyllum.

The known distribution of these four species is shown in text-fig. 1,
and the differences between their leaves in pi. VII, figs. 20-24. Maiden
and Baker (1895) described two other species in this group, viz.,
H. tesselatum and H. brevidecurrens — both from Murrumbo on the
upper Goulburn River, New South Wales. I have seen type materials
and, although their leaves are much longer and less appressed (fig. 23)
than in typical H. adnatum (fig. 22), with the heads considerably
larger (H. tesselatum also has more coarsely papillate achenes), the
differences from this species and one another do not seem to be specific.
Further investigation in the field may show that these are connected
with a variable H. adnatum by a series of intermediate forms.

104

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

Text Fig 1. — Distribution maps for Heliciirysum ,diosini folium (Vent.) Sweet,
H. adnatum (DC.) Benth., H. diotophyllum F. Muell. and H. tuclceri sp. nov. —
from material in Melbourne, Sydney and Brisbane Herbaria..

NOTES ON SOME AUSTRALIAN COMPOSITAE.

105

Chthonocephalus multiceps J. H. Willis; species nova ob capita
composita numerosa dissita a tribns aliis speciebus Chthonocephali
valde distincta; C. tomentellus (F. Muell.) Benth. liabitn
staturaque simillimus est, sed capitibus terminalibns solitariiss
floribns pluribus et pappo nullo differt.

Herba annua rosulata pluricaulis. Caules nigri, subfiliformes,
radiantes, subprostrati, usque ad 7 cm. longi, fragmenta tomenti
arachnoidei ferentes. Folia caulis extremitatem versus decrescentia ;
inferiora atque vetustiora usque ad 2-5 cm. longa, oblanceolata,
subglabrescentia ; superiora eonferta, 3-5 mm. longa, obovata, valde
lanigera pilis adpressis intrieatis basi infiatis 30 mic. latis, apiee
longe flagellatis. Capita composita inflorescentiae usque 40 per plantain,
axillaria atque terminalia, subdissita, sessilia, pallide flavescentia,
modice lanuginosa; summa maxima, congesta, pulviniformia, 5-12 mmu
diam. Capitula usque 40 in capitibus majoribus, 3-6 flora, receptaculo
spiculam brevissimam formanti. Involucri squamae usque ad 3 mm.
longae, dimorphae; paucae bracteae exteriores stipite angusto, praeter
basin laciniatae atque intricate lanuginosae ; bracteae interiores 4-7,
obovatae glabrae, hyalinae, manifeste areolatae, marginibus subintegrae.,
Flores 2-2*5 mm. longi, quisque ad basin a bractea lanata partim clausus
(bracteae subtendentes involucri squamis exterioribus simillimae) ;
corolla anguste infundibulifonnis, lobis 5 conspicuis omnino glabris.
Pappi squamae 5-7, quam corolla circiter triplo breviores, ad basin
connatae, laciniatae, supra pilis intrieatis ornatae, in floribus summis
breviores, paucilobati, glabri. Achaenia triquetra, obovoideo-turbinata
stylo-basi persistenti, 0-5-0-8 mm. longa, nubila, omnino minute papillosa~
Styli ramuli 0*5-1 mm. longi, ad apicem penicillati papillis longis
radiantibus. Antherae 1-1*5 mm. longae apiculis sterilibus conspicuis,
caudis longis gracilibus; filamenta perbrevia, prope medium corollae
tubum affixa. Pollinis granula 18-20 mic. diam., acriter echinulata. —
Plate VII, figs. 25~33.

Western Australia. — South-East Division (Euclonia) : Balladonia Homestead,
128 miles east of Norseman, on sandy soil against large granite slabs, with ephemeral
Helipterum spp., etc., 31st August, 1947, J. H. Willis, Grimwade Expedition

(HOLOTYPE in MEL, PARATYPE in PERTH, CLASTOTYPE in BRI.).

Both the author and Mr. C. A. Gardner, who examined the material
24/8/1948, at first considered this remarkable plant to be a species of
Gnephosis which it resembles outwardly ; but the presence of subtending
scales between the florets shows its affinity with Craspedia and
Chthonocephalus in the sub-tribe Angiantlieae. Members of the former
genus have long, erect peduncles, a general involucre, long, plumose
pappus scales, and a silky-hairy achene. The newly described plant
lacks these features, and conforms well to the present circumscription
of Chthonocephalus, making a fourth species in that genus. In its
growth form C. multiceps approximates to the west-coast (Western
Australia) species C. tomentellus, but in this the inflorescences are wholly
terminal on the woolly branches and the numerous florets are completely
devoid of any pappus. The only other species with a pappus is
C. pygmaeus — a rare, minute, stemless plant with narrowly linear leaves
— which could never be mistaken for the new one. C. multiceps is at
once recognizable by its very numerous compound heads (to 40) of
varying sizes, and it is only to be expected that it will prove to have
a wide distribution in the botanically little-known south-eastern part
of Western Australia.

106 PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

Senecio runcinifolius J. H. Willis, nomen novum.

Erechthites picridioides Sonder & F. Muell. in Linnaea 25: 523
(1852), non E. picridioides Turczaninow, 1851.

E. mixta sensu Benthami in FI. Aust. 3: 659 (1866), non (A.
Rich.) DC., 1837.

♦

South Australia. — Murray River at Moorundee near Blanehetown, Feb. 1851,
F. Mueller (holotype & paratypes in mel) • “Towards Spencer Gulf”,
Warburton.

Victoria. — -North-West: Between Cohuna and Kerang, May, 1944, B. V. Smith;
Berribee Tank, far NW. corner of State (in a. large claypan 2 piiles east), 31st
Aug., 1948, J. H. Willis.

New South Wales. — Far Western Plains: Junction of Murray and Darling
Rivers, 1889, Mrs. Molding. Western Plains: Warrego River, E. Betche.

The epithet of the new name (rendered necessary by preoccupation
of the name Erechthites picridioides) is in allusion to the remarkable,
retrorsely-lobed leaves of this plant that resemble those of a dandelion
( Taraxacum species). It would seem to be riparian and, in view of
the scanty material available in Australian herbaria, is apparently rare.
The original diagnosis, drawn up from poor “starved” specimens, gives
a completely inadequate picture of the species and an amended and
amplified description follows:

Herb, pale green, glabrescent, 20-40 cm. high (6 ins. in original
description), the young parts and leaf axils slightly cottony- woolly.
Leaves sessile, lanceolate acuminate, with up to 8 more or less reversed
sinuate-lobed segments on each side, to 12 cm. long and 3 cm. wide at
base of stems (1 in. and 3-4 lin. in original description) ; upper floral
leaves less toothed, with long, almost filiform tips. Panicle corymbose,
loose, up to 60-headed (6-8 in original description). Peduncles of
each head slender, up to twice the length of . involucre which is minutely
cottony, bracteolate at the base. Phyllaries of involucre 12-14, about 1
cm. long, linear, acute, glabrous in fruit. Corolla tube of central
hermaphrodite florets about 0-5 mm. diameter at base of the 5 lobes,
about 4 times wider than the tubular filiform female florets; lobes in
both with shortly papillose margins. Achene 2-5-3 mm. long, pallid,
slightly flattened (about 0-3 mm. wide), with about 9 costae that bear
short, erect, papillate hairs — without the attenuated beak ascribed to
it in the original diagnosis. Anthers about 1 mm. long, without basal
appendages. Style arms ligulate, about 0-33 mm. long, with swollen,
recurved apices, coarsely papillose on inner face. Pollen grains sharply
echinulate, 20-25 mic. diameter. — Plate VII, figs. 34-37.

All the collections cited are in Melbourne Herbarium, but Bentham
saw only the South Australian ones. It is astonishing he should have
confused E. picridioides Sonder & F. Muell. with E. mixta (A. Rich.)
DC. — a purplish mountain plant of entirely different aspect and floral
structure. JVM. Black (1929) perpetuated the confusion.

[Mr. A. O. Belcher (Michigan) is revising Erechthites Rafin., and many changes
are anticipated in the nomenclature of Australian species hitherto referred to this
genus.]

NOTES ON SOME AUSTRALIAN COMPOSITAE.

107

REFERENCES.

Black, J. M., 1929. Flora of South Australia 4: 610.

Ewart, A. J., 1930. Flora of Victoria, 1137.

Maiden, J. H., and Baker, R. T., 1895. Proc. Linn. Soc. N.S. Wales 10: 589, 590.

EXPLANATION OF PLATE VII.

Figs. 1-7, Helichrysum erioeephalum sp. nov. — 1, branch with inflorescence
(1 nat. size); 2, capitulum; 3, upper involucral bract; 4, central floret; 5, tip of
pappus bristle; 6, style arms; 7, anther.

Figs. 8-10, H. cordatum DC. (upper bract, central floret and pappus bristle — '
cf. figs. 3-5).

Figs 11-13. H. ramosum DC. (upper bract, central floret and pappus bristle —
cf. figs. 3-5).

Figs. 14-20, Helichrysum tuckeri sp. nov. — 14, branch with inflorescences
(| nat. size); 15, capitulum; 16, upper involucral bract; 17, central floret; 18, tip
of pappus bristle; 19, anther; 20, leaves (enlarged).

Figs. 21-22, enlarged leaves of H. diotophyllum F. Muell. and H. adnatum
(DC.) Benth. respectively. — cf. with fig. 20.

Figs. 23-24, leaves of H. tesselatum Maiden & Baker and H. diosmifolium
(Vent.) Sweet, respectively (f nat. size). — cf. figs. 20-22.

Figs. 25-33 Chthonocephalus multiceps sp. nov. — 25, plant (4 nat. size) ;
26, flagellate hairs on leaf surfaces; 27, hairless inner involucral bract of partial
head; 28, outer bract of partial head; 29, uppermost floret, with hairy subtending
scale and reduced pappus; 30, lower floret, with normal pappus; 31, triquetrous
achene; 32, style; 33, anther (inserted on corolla tube).

Figs. 34-37, Senecio runcinif dlius nom. nov. — 34, branch with an inflorescence
(4 nat. size); 35, outer female and central hermaphrodite florets; 36, style arms;
37, anther.

[Except in figs. 1, 14, 25 and 34, a scale in millimetres accompanies each
drawing; the scale for figs. 23-24 is in centimetres.]

Proc. Roy. Soc. Q’land, Yol. LXII., No. 11.

Plate YII.

Some Australian Compositae.

Vol. LXIL, No. 12.

10!)

WHITEOCHLOA, A NEW GENUS OF GRASSES
FROM THE NORTHERN TERRITORY
OF AUSTRALIA.

By C. E. Hubbard, Royal Botanic Gardens, Kew, England.

(. Issued separately, 22nd August, 1952.)

SUMMARY.

WMteochloa, a new genus of Gramineae, tribe Paniceae, is described.

The tribe Paniceae of the family Gramineae is exceptionally well-
developed in the warmer parts of Australia, no fewer than thirty-nine
genera now being recorded as native or naturalized and well-established,
a very considerable increase on the thirteen described by Bentham in
the Flora Australiensis (1878). This trebling of the number of genera
is due mainly to a more uniform conception of these taxa than that
adopted by Bentham, the genus being now restricted to species very
similar in the structure and arrangement of their spikelets, and
presenting a distinctive facies. Thus D. K. Hughes (1923), in her
revision of the genus Panicum of the Flora Australiensis accepted the
genera defined by A. Chase and by Stapf as a result of their studies
of American and African Paniceae respectively, and divided the species
of Panicum sensu Bentham among fourteen genera. Further sub-
division has been carried out by S. T. Blake in segregating his new
genera Zygochloa from Spinifex and Ancistrachne from Panicum, and
by Pilger in removing Pseudor aphis from Chamaer aphis. On the other
hand, several additions to the Australian genera of Paniceae are due to
more intensive collecting in botanically unexplored areas; they include
the genera Calyptochloa, Cleistochloa, Dimorphochloa, Homopholis,
Ottochloa, Pseudochaetochloa and TJranthoecium. A few exotic genera
introduced as fodder plants, such as Axonopus, Melinis and
Bhynchelytrum, have become established in the warmer parts of the
Commonwealth.

The most important characters used in distinguishing the genera
of the Paniceae are to be found in those modifications of the scales of
the spikelet which ensure greater protection or more widespread
dispersal of the seed, in those produced by lateral or dorsal pressure
during the development of the spikelet, in the degree of development
of the lower floret, in the arrangement of the spikelets in the inflorescence
and in the form of the latter. Among the Australian species referred
to the genus Paspalidium Stapf by Hughes, is one, P. semitonsum
(F. Muell. ex Benth.) Hughes (l.c. 317), which differs from all other
members of the genus in so many of these diagnostic characters that
it must be separated as a distinct genus. The name Whiteochloa is

110

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

proposed for this new genus. The distinctions between the two genera

are set out in the following table :

Paspalidium.

1. Spikelets plano-convex or slightly
dorsally compressed.

2. Upper floret sessile on the rhachilla.

3. Lower glume ab axial ; upper glume
adjacent to the axis on which the
spikelet is borne.

4. Upper glume glabrous.

5. Lower lemma usually flat or slightly
depressed on the back.

6. Upper floret about as long as the
lower.

7. Axes of the inflorescence each ter-
minated by a bristle or blunt naked
tip.

Whiteochloa.

1. Spikelets slightly laterally com-
pressed.

2. Upper floret borne on a very short
rhachilla-internode.

3. Lower glume adaxial, adjacent to the

axis on which the spikelet is borne.

4. Upper glume bearing a row of stiff
tubercle-based hairs on each nerve.

5. Lower lemma very convex on the
back except for a very narrow
shallow median translucent longi-
tudinal groove.

6. Upper floret shorter than the lower.

7. Axes of the inflorescence each ter-
minated by a spikelet.

Judging from the orientation of its spikelets and by their general
structure, Whiteochloa is more closely related to Pseudechinolaena Stapf
than to Paspalidium. This genus of forest grasses has been recorded
from Papua and may occur in the rain forests of northern Queensland.
It comprises two species, Pseudechinolaena polystachya (H.B.K.) Stapf,
with a wide area of distribution in tropical America, Africa and Asia,

and P. perrieri A. Camus, which is
genera may be distinguished by the
Pseudechinolaena.

1. Glumes equal, herbaceous, or the lower
slightly shorter, rarely only two-
thirds the length of the spikelet.

2. Upper glume with more or less
translucent spots between the nerves,
and usually with a straight or
eventually hooked many-celled bristle-
like outgrowth from the centre of
each spot.

3. Lower lemma convex on the back.

4. Ligules membranous.

5. Leaf-blades lanceolate to ovate,
short.

confined to Madagascar. The two
following characters : —

Whiteochloa.

1. Glumes unequal, membranous, the
lower one-third to half the length of
the spikelet.

2. Upper glume bearing a row of stiff
tubercle-based unicellular white hairs
along each nerve.

3. Lower lemma convex on the back
except for a narrow median groove.

4. Ligule reduced to a ciliate rim.

5. Leaf-blades narrowly linear,

elongated.

It is very probable that the two genera differ much in habit, the
species of Pseudechinolaena being trailing annuals or perennials,
branching and rooting from the prostrate many-noded base, and with
very slender ascending leafy culms. Unfortunately the type-material
of Whiteochloa lacks the base, but it has the appearance of an erect
or suberect annual, with simple few-noded culms.

The genus Ancistrachne S. T. Blake resembles Whiteochloa in some
respects, but may be readily separated by its rigid woody branched
culms, the slightly dorsally compressed abaxial spikelets, 9-11-nerved
upper glume, barren lower floret, 7-9-nerved lower lemma and by the
flattened margins of the upper lemma. The genus Brachiaria, which

WHITEOCHLOA, A NEW GENUS OF GRASSES.

Ill

agrees with Whiteochloa in possessing adaxial spikelets, may be
distinguished by its terete or dorsally compressed spikelets, sessile upper
floret, absence of tubercle-based hairs on the nerves of the upper glume,
and by the thin median groove of the coriaceus lower lemma.

Generic Description.

Whiteochloa C. E. Hubbard, genus novum, affine Pseudechinolaenae
Stapf, sed glumis membranaceis, gluma inferiore late ovata usque
dimidiam partem spiculae aequante, gluma superiore secus nervos pilis
brevibus rigidiusculis e tuberculis minutis ortis ciliata, lemmate infero
dorso longitudinaliter sulcato, ligula ad seriem ciliorum redacta, laminis
foliorum anguste linearibus differt.

Spiculae asymmetricae, a laterae visae semi-ovatae vel semi-elliptico-
ovatae, a dorso visae anguste ellipticae et acutae, exaristatae, leviter
lateraliter compressae, contiguae vel imbricatae, adaxiales, breviter et
inaequaliter pedicellatae, demum totae a pedicellis persistentibus
disarticulantes, solitariae vel binae, in ramis et ramulis secundis
panicularum erectarum linearium vel lanceolatarum dispositae; rami
gracillimi, triquetri, solitarii, inferne ramulos breves appressos
paucispiculatos gerentes; rhachilla inter anthoecia internodio gracillimo
brevissimo glabro praedita. Anthoecia duo, dissimilia ; inferum
masculum ; superum hermaphroditum, infero brevius. Glumae
inaequales, dissimiles ; inferior usque dimidiam partem spiculae aequans,
late ovata, obtusa vel acuta, mucronulata, membranacea, 3-5-nervis ;
superior spiculae aequilonga vel fere aequilonga, cymbiformis, dorso
valde convexa, explanata anguste ovata, acute acuminata, membranacea,
5-7-nervis, nervis pilis brevibus patulis rigidiusculis e tuberculis minutis
ortis ciliata. Anthoecium inferum: lemma spiculae aequilongum vel
fere aequilongum, dorso longitudinaliter anguste sulcatum, ceterum
convexum, sulco translucente excepto coriaceum, elliptico-oblongum,
subacutum vel obtusum, tenuiter 5-nerve, glabrum vel fere glabrum,
palea anguste elliptica vel oblonga, lemmati aequilonga vel eo paullo
longior, obtusa, bicarinata, circa carinas anguste alatas indurata, ceterum
tenue membranacea. Anthoecium superum a latere visum semi-elliptico-
ovatum, a dorso visum anguste ellipticum et acutum : lemma apiculatum,
dorso valde convexum, marginibus involutis angustis firmis, demum
crustaceum, 'tenuiter 5-nerve, tenuiter transverse rugulosum; palea
lemmati aequilonga, dorso plana, Crustacea, 2-nervis. Lodiculae 2, late
oblongae, truncato-emarginatae, glabrae. Stamina tria ; antherae anguste
oblongae. Ovarium glabrum; styli liberi, terminales; stigmata breviter
plumosa. Caryopsis a dorso visa elliptica, dorso compressa, plano-
convexa; scutellum circiter dimidiam partem caryopseos aequans; hilum
basale, ellipticum. — Culmi graciles, erecti, simpliees; foliorum vaginae
anguste lineares, planae ; ligulae ad seriem ciliorum redactae ; paniculae
elongatae.

Species unica, Australiae tropicae incola.

Whiteochloa semitonsa ( F . Muell. ex Benth.) C. E. Hubbard, comb. nov.

Panicum semitonsum F. Muell. ex Benth. FI. Austral. 7 : 483
(1878) ; Ewart & Davies, FI. North. Territ. 39 (1917).

Paspalidium semitonsum (F. Muell. ex Benth.) Hughes in Kew
Bull. 1923: 317 (1923).

112 PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

Gramen annum (?), basis ignotum, circiter 60 cm. altum. Culmi
glabri, laeves. Foliorum vaginae marginibus apicem versus ciliolatae,
ceterum glabrae; laminae usque 12 cm. (vel ultra) longae, 3.5 mm.
latae, supra minute scaberulae, glabrae. Inflorescentia 10-20 cm. longa,
usque 5 cm. lata; rami inferiores usque 7 cm. longi; pedicelli 0-3-2
mm. longi. Spiculae 3-5-4-2 mm. longae; internodium rhachillae usque
0-5 mm. longum; gluma inferior 1-5-2 mm. longa, nervis minute
hispidula; anthoecium superum 2-2-2-5 mm. longum; antherae 1-3-1 -5
mm. longae.

Northern Territory: Victoria River, Elsey! (Herb. Kew.).

In addition to the above, Bentham (l.c.) also cites a specimen
collected by F. Mueller at Providence Hill, north of the mouth of the
Victoria River, at about 14° 30' S and 129° 30' E. This material was
probably returned to the Melbourne Herbarium. Both J. R. Elsey ’s
and F. Mueller’s specimens were gathered on A. C. Gregory’s Northern
Australian Expedition of 1855-56, on which the former served as
surgeon and naturalist and the latter as botanist.

The name Whiteochloa is given in memory of Cyril Tenison White,
one of Australia’s most illustrious botanists. He is remembered by the
writer with gratitude and affection as a very good friend, who by his
generous help, kindly advice and encouragement, and perfect companion-
ship on numerous botanical excursions, made most memorable a year
spent at the Brisbane Herbarium and elsewhere in Queensland in
1930-31.

REFERENCES.

Bentham, G., 1878. Flora Australiensis 7: 452-3.

Hughes, D. K., 1923. The genus Panicum of the Flora Australiensis. Kew Bull.
1923: 305-332.

The Royal Society of Queensland.

Report of the Council for 1949.

To the Members of the Royal Society of Queensland .

Your Council has pleasure in submitting the Annual Report of
the Society for the year 1949.

At Ordinary Meetings throughout the year five addresses were
given, one film was shown, and one exhibit evening held; while on one
occasion a report was made on the Science House project. The Annual
Memorial Lecture, held this year in honour of Mr. F. M. Bailey, was
delivered by Mr. C. T. White. Eight original papers were accepted
for publication in the Proceedings.

An approach was made to the Premier and Chief Secretary’s
Department for an increase in the maximum yearly subsidy payable
on publication, and an increase from £150 to £200 was granted.

To overcome the delay in publication of the Proceedings, the
Council has decided, in agreement with the Government Printing Office,
to have the Volume for 1949 printed by an outside firm, in which case
the Government subsidy will still be available.

The Society has obtained sole occupancy of the room in which the
Library is housed, and additional shelving has been constructed. The
Library is now in quite good order, but still requires cataloguing.

There are 5 honorary life members, 9 life members, 3 corresponding
members, 234 ordinary members and 1 associate member in the Society.
This year the Society has lost 1 member by death and 10 by resigna-
tion; 22 ordinary members and 1 associate member have been elected,
and 1 member has been readmitted.

Attendance at Council Meetings was as follows : — D. Hill, 10 ;
H. C. Webster, 9; M. F. Hickey, 7 ; M. I. R. Scott, 10; D. F. Sandars,
9; B. Baird, 8; S. T. Blake, 9; G. Mack, 7; O. A. Jones, 9; A. L.
Reimann, 7 ; J. H. Simmonds, 5 ; L. J. H. Teakle, 5 ; E. M. Shepherd, 8.

DOROTHY HILL, President.

Margaret I. R. Scott. Hon. Secretary.

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Hon. Auditor. DOROTHEA SANDARS, Hon. Treasurer.

ABSTRACT OF PROCEEDINGS.

VII.

Abstract of Proceedings, 27th March, 1950.

The Annual General Meeting of the Society was held in the Geology
Department of the University on Monday, 27th March, with the
President (Dr. Dorothy Hill) in the chair. About forty-five members
and friends were present. Apologies were received from His Excellency
the Governor, Prof. Bostock, Dr. Nye and Mr. Longman. The Minutes
of the last Annual General Meeting were read and confirmed. The
Annual Report was adopted and the Balance-sheet received. The
following were nominated for membership: — Mr. F. W. Berrill,
Dr. Ernest Singer, Mr. R. F. Isbell, Mr. G. A. Wyatt, Mr. I. F. Fergus
and Mr. J. D. Hughes for Ordinary Membership, and Miss B. Howard,
Miss M. Patey, Miss J. W. Herbert, Miss D. J. Matthews and Miss H. M.
Simmons for Associate Membership.

The following officers were elected for 1950 : —

President: Prof. M. F. Hickey.

Vice-President : Prof. H. J. G. Hines.

Hon. Secretary : Miss M. I. R. Scott.

Hon. Treasurer : Miss D. F. Sandars.

Librarian: Mr. F. S. Colliver.

Editors: Mr. S. T. Blake, Mr. G. Mack.

Members of Council : Dr. I. M. Mackerras, Prof. A. L. Reimann,
Mr. J. H. Simmonds, Prof. L. J. II. Teakle, Prof. II. C.
Webster.

Hon. Auditor : Mr. L. P. Herdsman.

The Presidential Address, entitled “The Earliest Corals,” was
delivered by Dr. Dorothy Hill. A vote of thanks was moved by Prof.
F. W. Whitehouse, seconded by Prof. W. II. Bryan and carried by
acclamation.

Abstract of Proceedings, 24th April, 1950.

The Ordinary Monthly Meeting of the Society was held in the
Geology Department of the University on Monday, 24th April, with
the President (Professor M. F. Hickey) in the chair. About sixty
members and friends were present. The minutes of the previous
meeting were confirmed. The following members were elected : —
Ordinary Members — Mr. F. W. Berrill, Dr. Ernest Singer, Mr. R. F.
Isbell, Mr. G. A. Wyatt, Mr. I. F. Fergus, Mr. J. D. Hughes, Miss D. J.
Matthews and Miss H. M. Simmons; Associate Members — Miss B.
Howard, Miss M. Patey and Miss J. W. Herbert. Miss Matthews
and Miss Simmons, who had been nominated as Associate Members
in error, had consented to stand for election as Ordinary Members.
The following nominations for ordinary membership were received: —
Mr. W. G. Burns, Mr. M. Strohfeldt, Mr. J. A. Thomas, Mr. D. Sinclair,
and Mr. M. Crawfoot. The Clarke Memorial Medal for 1950 was
presented on behalf of the Council of the Royal Society of New South
Wales to Dr. Ian M. Mackerras.

Dr. Owen Jones gave a lecture on “The Use of Seismographs in
the Detection of Cyclones. ’ ’ The method utilizes three sensitive seismo-
graphs set up at the apices of an equilateral triangle. The bearing of

VIII. ABSTRACT OF PROCEEDINGS.

the storm-centre is calculated from the differences in time of arrival
of an identical microseismic wave at the three instruments. The
position of the centre is fixed by intersection of bearings from two or
more stations. Professor W. H. Bryan was the first (in 1938) to note
the correlation of microseisms and hurricanes. Since then work at
the University of Queensland Seismological Station has shown: —

(1) That, even with our present instruments, which are not as
sensitive as those used by the U.S. Navy, we can detect
hurricanes.

(2) That, in some cases at least, we can detect their presence
before they can be detected by the methods now in use by
the Weather Bureau.

(3) That we can distinguish between malevolent and beneficent
cyclones.

(4) That the use of microseisms would be a valuable supplement
to the present methods of locating and following the course
of hurricanes.

We cannot, however, locate them, that is, determine the bearing
of the centre. For that we need more seismographs of an even more
sensitive type. To equip four stations in Queensland would cost about
£28,000 and about £500 per year in upkeep.

Abstract of Proceedings, 22nd May, 1950.

The Ordinary Monthly Meeting of the Society was held in the
Geology Department of the University on Monday, 22nd May, with
the President (Dr. M. F. Hickey) in the chair. About thirty-five
members and friends were present. The minutes of the previous
meeting were confirmed. The following were elected to Ordinary
Membership : — Mr. W. G. Burns, Mr. M. Strohfeldt, Mr. J. A. Thomas,
Mr. D. Sinclair and Mr. M. Crawfoot.

Dr. E. J. Reye exhibited some live plankton from Moreton Bay.

Three short addresses were given by Dr. M. F. Hickey, Professor
A. L. Reimann and Miss D. Sandars, respectively.

In discussing “Some Aspects of Congenital Abnormalities, ’ ’
Dr. Hickey gave a brief illustrated account of the development of the
fertilized human ovum to the stage of the three-layered embryonic
disc, and indicated the various ways in which it was thought that
identical twins might develop during these early stages and how certain
of the forms of so-called “Siamese twins” probably developed. He
indicated the difficulty of establishing ultimate causes, but pointed out
that the work of Streeter, Douglas Murphy and others leads to the
general conclusion that gross human congenital malformations arise
solely from influences which affect the germ cells prior to fertilization.
But in the case of fishes, amphibia and birds, it appeared that mal-
formations could be produced in the developing embryo by alterations
of the environment — chemical, physical and thermal.

Professor Reimann spoke on electron gases. The properties of
ordinary and electron gases were compared and contrasted, with special
reference to (a) condensation, evaporation and latent heat; (b) winds;

ABSTRACT OF PROCEEDINGS.

IX.

and (c) the distribution of thermal velocities. Under (a) it was
pointed out that as in the case of the molecules of ordinary gases, so also
electrons may be obtained in the free state by evaporating them from a
condensed phase, e.g., from metals (thermionic emission), and that a
latent heat of evaporation has to be supplied to bring this about, this
being related to the thermionic work function. Under (5) it was shown
that the operation of electronic devices such as radio valves, X-ray tubes,
cathode-ray tubes and electron microscopes depends on the control of
electron “winds” derived from a thermionic cathode and directed, by
appropriate electric or magnetic fields, to an anode, in which latter
they are condensed. Under item (c), it was shown how, in principle,
the range of gas thermometry, on which our standard scale of tem-
perature is based, might be extended far beyond the upper practical
limit for ordinary gases, viz., about 1,500 deg. C., by using an electron
gas in place of an ordinary gas such as nitrogen. Electron-gas ther-
mometry would have to be based on the theoretical relation between
velocity distribution of emission and temperature, the former being
explored by the application of various retarding potentials to the anode.
In this way gas thermometry could be extended to near the melting
point of tungsten (about 3,400 deg. C.).

Miss Sandars spoke on the Great Barrier Reef Committee’s pro-
posal for a marine biological station. She said that the committee has
decided to establish a marine biological station on the Reef for research
into reef biology and associated problems, for training young research
workers, and for research into marine economic problems. It could
also serve as a centre for other scientific investigations. Heron Island
(Capricorn Group) has been chosen as the site for the station, and
draft plans of the building and estimates of costs have been prepared.
The proposals have been approved by A.N.Z.A.A.S., A.N.R.C.,

C.S.I.R.O., The Pacific Science Association, the Royal Society and the
Royal Geographical Society. The estimated capital cost is £5,000 for
building and £500 for basic equipment. Already £2,468 14s. Id. has
been received, including £1,000 stg. from the M. T. Browne fund of the
Royal Society, £500 from the Great Barrier Reef Committee, £517 9s. Id.
from the Goddard Memorial Fund, and £201 5s. from other organisa-
tions and private donations. The Government has accepted gifts to
the fund as rebatable for taxation purposes, and the Committee is now
seeking to augment it substantially by individual donations. The
support of the Royal Society of Queensland was expressed in the
following motion, proposed, seconded and carried unanimously : —

“That the Royal Society of Queensland commends the
proposal of the Great Barrier Reef Committee to establish a
marine biological station on the Reef and offers its influence in
support of the project.”

Abstract of Proceedings, 26th June, 1950.

The Ordinary Monthly Meeting of the Society was held in the
Geology Department of the University on Monday, 26th June, with the
President (Dr. M. F. Hickey) in the chair. About forty members and
friends were present. The minutes of the previous meeting were con-
firmed. The following were nominated for Membership : — Mr. J. P.
Webb and Miss Rona E. Stewart — Ordinary Members; Mr. J. S.

ABSTRACT OF PROCEEDINGS.

Derrington, Mr. C. W. Siller, Mr. I. R. McLeod, Mr. J. B. Jones,
Mr. A. R. Lloyd, Mr. J. P. Stephenson and Mr. K. G. Smith —
Associate Members.

Dr. E. Singer gave an address entitled “Four Years in China.”

Abstract of Proceedings, 31st July, 1950.

The Ordinary Monthly Meeting of the Society was held in the
Geology Department of the University on Monday, 31st July, with the
President (Dr. M. F. Hickey) in the chair. About forty members
and friends were present. The minutes of the previous meeting were
confirmed. The following were elected to membership: — Mr. J. P.
Webb, Miss R. E. Stewart — Ordinary Members; Mr. J. S. Derrington,
Mr. C. W. Siller, Mr. I. R. McLeod, Mr. J. B. Jones, Mr. A. R. Lloyd,
Mr. J. P. Stephenson, Mr. K. G. Smith — Associate Members. The
following were nominated for Associate Membership : — Miss J. von
Alpen, Miss C. Goldsmid, Mr. W. R. Dowd.

The President made the following announcement regarding the
death of Dr. Gustave Athol Waterhouse. 6 4 1 regret to announce that
Dr. Gustave Athol Waterhouse died in Sydney on the 29th July. He
was not a member of this Society; but he was well known to many
members as an authority on Australasian Lepidoptera, and he was for
many years prominent in the councils of our sister Royal Society in
New South Wales, as well as of the Linnaean Society, the Australian
National Research Council, and other scientific organisations. Our
sympathy is extended to his family.”

The Librarian reported that 76 additional volumes and parts had
been added to the library since the last meeting. Also a new exchange
has been established for the Comptes Rendus d’Academie des Sciences,
Bulgaria.

The following paper was presented: — “Aphistomyia collini Bezzi
(Diptera, Blepharoceridae) in North Queensland” by I. M. Mackerras
and M. J. Mackerras.

Professor W. Stephenson gave an address entitled ‘‘Preliminary
Observations upon the Evolution of Phosphates from Estuarine Muds.”
Some of the difficulties of this type of investigation, even when under-
taken under laboratory conditions, were detailed. Muds sometimes
absorb phosphate, and sometimes release it, and the presence of animals
in the mud may increase absorption or initiate release. Evolution of
phosphate from surface deposits of mud proceeds rapidly under con-
ditions of oxygen lack, and as previously shown (1949), phosphate
may be evolved from mud and mud filtrates merely by agitation.

Abstract of Proceedings, 4th September, 1950.

The Ordinary Monthly Meeting of the Society was held in the
Geology Department of the University on Monday, 4th September,
with the President (Dr. M. F. Hickey) in the chair. About thirty-six
members and friends were present. The minutes of the previous
meeting were confirmed. Miss C. Goldsmid and Miss J. von Alpen
were elected to Associate Membership.

ABSTRACT OF PROCEEDINGS.

XI.

A film entitled “Smoke Streams” was shown, a commentary being
given by Mr. G. Birkbeck.

Professor M. Shaw exhibited partially made experimental models
of a cutting-tool dynamometer for measuring steady and also transient
cutting forces, and a profilometer for measuring surface finish.

Professor W. H. Bryan exhibited a fossilized branch of a coniferous
tree in the shape of a hollow mould 27 cm. in length and 5 cm. in
breadth from the lower (non-welded) part of the Brisbane Tuffs as
developed at Butterfield road, near the Brisbane General Hospital.
The fossil was from approximately the same horizon as that in which
numerous fossil trees have been found, the petrified wood of which
is, in most cases, closely comparable with that of modern conifers such
as Araucaria.

Professor D. A. Herbert exhibited (a) portions of the trunks of
two rain-forest saplings that had covered encircling vines with callus
and continued to grow with the vine stems apparently entering below
and emerging further up, (&) a trunk of Myrtus hillii which had
grown over a vine lying in a fork so that the vine stem appeared to
be growing through the wood, and (c) some dressed planks of hoop
pine ( Araucaria cunninghamii) with vines deeply embedded and
covered with several inches of wood. In the hoop pine specimens the
result was curious in that knots of dicotyledonous wood were in the
gymnosperm timber.

Dr. M. F. Hickey exhibited some micro-photographs in colour
of sections of tissue showing the large multi-nucleated cells usually
called Osteoclasts (various magnifications up to x 850). The slides
illustrated the great variety of shape of these cells, their situation as
usually described, i.e., at areas of- bone absorption, but also apparently
in areas where bone deposition was in its earliest stage, and some in
sites apparently remote from bone formation. Some of the cells showed
marked pseudopodial processes. The speaker referred to a review of
the problem of the Osteoclast by Hancox containing an extensive list
of references in Biol. Rev., Yol. 24, pp. 448-471, October 1949.

Abstract of Proceedings, 25th September, 1950.

The Ordinary Monthly meeting of the Society was held in the
Geology Department of the University on Monday, 25th September,
with the President (Dr. M. F. Hickey) in the chair. About eighty
members and friends were present. The minutes of the previous
meeting were confirmed. Mr. R. H. Greenwood was nominated for
Ordinary Membership. The Librarian reported the addition of about
ninety volumes and parts to the Library since the last meeting.
Mr. L. C. Ball was elected to Honorary Life Membership. Mr. W. P.
Dowd was elected to Associate Membership.

Three films were shown : — -

( 1 ) “In All Weathers. ’ ’

(2) “Wonders of the Deep.”

(3) A film of marsupial mice.

XII.

ABSTRACT OF PROCEEDINGS.

Abstract of Proceedings, 30th October, 1951.

The Ordinary Monthly Meeting of the Society was held in the
Geology Department of the University on Monday, 30th October, with
the President (Dr. M. F. Hickey) in the chair. About sixty members
and friends were present. The minutes of the previous meeting were
confirmed. Mr. R. H. Greenwood was elected to Ordinary Membership.
Dr. G. C. Kenny and Mr. C. G. Ludford were nominated for Ordinary
Membership. The Librarian reported that 89 volumes and parts have
been added to the Library since the last meeting ; also two new exchanges,
Die Erde and Ann. Fac. des Sciences de Marseille, have been established.

Mr. F. S. Colliver exhibited, on behalf of the Geology Department,
parti-coloured tourmaline crystals from Arizona.

Professor F. W. Whitehouse exhibited a late Tertiary basalt collected
by Mr. Ogilvie north of Hughenden, showing curious radial rosette
structures only doubtfully of spherulitic origin.

Professor J. T. Wilson, of Toronto, gave an address entitled “The
Growth of a Continent. * 9

Abstract of Proceedings, 27th November, 1950.

The Ordinary Monthly Meeting of the Society was held in the
Geology Department of the University on Monday, 27th November, with
the President, Associate Professor M. F. Hickey, in the chair. About
thirty members and friends were present. The minutes of the previous
meeting were confirmed. Dr. G. C. Kenny and Mr. C. G. Ludford
were elected to Ordinary Membership. Professor T. K. Ewer and
Professor F. T. M. White were nominated for Ordinary Membership.
The Librarian reported that 122 volumes and parts had been added to
the Library; also new exchanges had been established for the Pakistan
Journal of Science, the Records of the South Australian Museum, and
Zoological Papers of Victoria College, New Zealand; in addition,
holdings of the Ohio Journal of Science and the Royal Society of
Tasmania have been completed.

Miss D. F. Sandars exhibited the contents of a bandicoot’s
stomach.

Professor W. H. Bryan exhibited specimens of turquoise collected
by Dr. 0. A. Jones and himself from a cutting in Adelaide street,
in the heart of the city of Brisbane. (This is probably the same
locality as that recorded by Major Sankey many years ago.) He
pointed out that turquoise and wavellite have also been rediscovered
at Wilston Hill and at Stafford, but Sankey ’s locality at Victoria
Park has not been found. Mr. A. K. Denmead commented on the
exhibit.

Professor M. Shaw gave an address entitled “Graticules and their
Production.” In his opening remarks he gave the reasons for choosing
a certain design of graticule and the thickness of the lines in the pattern.
For example, a telescope sight for a jungle rifle must have a very
thick line to stand out against the confused background, whereas a
theodolite may have a very fine line of about 0-0001 in. in width.
Various types of graticules were shown and explained both by means of

ABSTRACT OF PROCEEDINGS.

XIII.

illustrations on the screen and also by the graticules themselves which
were on show. The speaker then went on to describe the methods used
in the manufacture of graticules of both the photographic and the
ruled and etched type. He spent some time explaining the instruments
which had been developed in the University of Melbourne, and illus-
trated his remarks by an interesting “Heath Robinson ” wooden model
which demonstrated in a very simple manner all the principles involved
in the extremely accurate prototype. Both the line-ruling and the
pantograph machines were explained. A full discussion followed which
was amplified by the remarks elicited as a result of a close examination
of the exhibit by the audience. A full account of this work is to be
obtained in the Proceedings of the Institution of Mechanical Engineers
160: 145 (1949).

XlV.

ALTERATIONS TO MEMBERSHIP.

Berrill, F. W.

Burns, W. G.

Crawfoot, A.

Fergus, I. F.
Greenwood, R. H.
Hughes, J. D.

Isbell, R. F.

Kenny, Dr. G. C.
Ludford, C. G.

Matthews, Miss D. J.
Simmons, Miss H. M.
Singer, Dr. E.

Stewart, Miss R. E.

Strohfeldt, M.

Thomas, J. A.

Webb. J. P.

Wyatt, G. A.

Derrington, J. S.
Dowd, W. R.
Goldsmid, Miss C.
Herbert, Miss J. W.
Howard, Miss B.
Jones, J. B.

Lloyd, A. R.

McLeod, I. R.

Patey, Miss M.

Siller, C. W.

Smith, K. G.
Stephenson, J. P.
von Alpen, Miss J. . .

New Honorary Life Member.

Ball, L. C.

New Ordinary Members.

Department of Agriculture and Stock, Nambour.

Geology Department, University, Brisbane.

Department of Agriculture and Stock, Brisbane.

Geography Department, University, Brisbane.

Department of Agriculture and Stock, Brisbane.

Geology Department, University, Brisbane.

Anatomy Department, University, Herston.

Queensland Institute of Medical Research, Herston road,
Brisbane.

Botany Department, University, Brisbane.

Ivy street, Indooroopilly.

Queensland Institute of Medical Research, Herston road,
Brisbane.

Radio Physics Laboratory University Grounds, City road,
Chippendale, N.S.W.

Physics Department, University, Brisbane.

Geology Department, University, Brisbane.

Department of Agriculture and Stock, Brisbane.

New Associate Members.

423 Milton road, Auchenflower.

Coronation drive, Auchenflower.

30 Eblin drive, Hamilton.

Botany Department, University, Brisbane.

Physiology Department, University, Brisbane.

Geology Department, University, Brisbane.

Physiology Department, University, Brisbane.

Geology Department, University, Brisbane.

Biochemistry Department, University, Brisbane.

Resignations.

Bosworth, F. O. Machin, W. F.

Gipps, R. de Y. Noyes, Miss M.

Jones, W. M. Tuffley, Mrs. A. M.

Lee, Dr. D. K.

Deaths.

White, C. T.

Henderson J. B.

A. H. Tucker, Governihent Printer, Brisbane.

GUIDE FOR THE PREPARATION OF SYNOPSES

1. PURPOSE.

It is desirable that each paper be accompanied by a synopsis preferably
appearing at the beginning. This synopsis is not part of the paper; it is intended
to convey briefly the content of the paper, to draw attention to all new information
and to the main conclusions. It should be factual.

2. STYLE OF WRITING.

It should be presumed that the reader has some knowledge of the subject
bul} has not read the paper. The synopsis should therefore be intelligible in itself
without reference to the paper, for example it should not cite sections or illustra-
tions by their numerical references in the text.

3. CONTENT.

It is sometimes valuable to indicate the treatment of the subject by such
words as: brief, exhaustive, theoretical, etc.

The synopsis should indicate newly observed facts, conclusions of an experiment
or argument and, if possible, the essential parts of any new theory, treatment,
apparatus, technique, etc.

It should contain the names of any new compound, mineral, species, etc., and
any new numerical data, such as physical constants; if this is not possible; it should
draw attention to them. It is important to refer to new items and observations,
teven though some are incidental to the main purpose of the paper; suchj information
may otherwise be hidden though it is often very useful.

When giving experimental results the synopsis should indicate the methods
used; for new methods the basic principle, range of operation and degree of
accuracy should be given.

4. DETAIL OF LAYOUT.

It is impossible to recommend a standard length for a synopsis. It should,
however, be concise and should not normally exceed 100 words.

If it is necessary to refer to earlier work in the summary, the reference should
always be given in the same manner as in the text. Otherwise references should
be left out.

CONTENTS

—

Vol. LXII.

No. 1. — The Ordovician Corals. By Dorothy Hill, D.Sc., Ph.D. (Issued

Pages.

separately, 15th October, 1951)

1-28

No. 2. — Apistomyia collini Bezzi (Diptera, Blepharoceridae) in
North Queensland. By I. M. Mackerras and M. J.
Mackerras. (Issued separately, 5th November, 1951)

29-32;

Technical Notes — Bandicoot Food. By Dorothea F. Sandars ...

C. T. WHITE MEMORIAL SUPPLEMENT.

No. 3. — Cyril Tenison White, 1890-1950. (Issued separately,

15th August, 1952) . . . . . . . . . .

35-48

No. 4. — Reductions in Elaeocarpus. By E. D. Merrill. (Issued
separately, 15th August, 1952)

49-56

No. 5. — Vegetative Habit in the Genus Eulophia (Orchidaceae).

By B. E. Holttum. (Issued separately, 15th August, 1952)

57-60

No. 6. — Rheophytes. By C. G. G. J. van Steenis. (Issued separately,
15th August, 1952)

61-68

No. 7. — PSEUDORAPHIS SPINESCENS (R.Br.) N. COMB., AND SOME
Records of New South Wales Grasses. By Joyce W.
Vickery. (Issued separately, 15th August, 1952) . .

69-72 |

No. 8. — The Significance of the Mallee Habit in Eucalyptus. By
N. T. Burbidge. (Issued separately, 29th August, 1952)

73-78 -

No. 9. — Opisthiolepis, a New Genus of Proteaceae from Queens-
land. By L. S. Smith. (Issued separately, 29th August
1952)

79-82

No. 10.— The Identification and Distribution of some Cyperaceae
and Gramineae, Chiefly from Australia. By S. T. Blake.
(Issued separately, 29th August, 1952)

83-100

No. 11. — Notes on Some Australian Compositae. By J. H. Willis.

(Issued separately, 22nd August, 1952)

101-108

No. 12. — Whiteochloa, a New Genus of Grasses from the Northern
Territory of Australia. By C. E. Hubbard. (Issued
separately, 22nd August, 1952)

109-112

Report of Council

V-VI

Abstract of Proceedings

VII-XIII

Changes in Membership

XIV

PROCEEDINGS

OF THE

ROYAL SOCIETY

QUEENSLAND

FOR 1951

VOL. LXIII.

ISSUED 17th AUGUST, 1953

PRICE: TWENTY-FIVE SHILLINGS.

Printed for the Society
by

A. H. TUCKER, Government Printer, Brisbane.

The Royal Society of Qyeensland.

Patron :

HIS EXCELLENCY LIETJT. -GENERAL SIR JOHN D. LAVARACK, C.B.,
C.M.G., D.S.O., C. de G., K.B.E.

OFFICERS, 1951.

President :

Associate Professor H. J. G. HINES, B.Sc.

Vice-Presidents :

M. E. HICKEY, M.A., M.B., B.S.
I. M. MACKERRAS, P.R.A.C.P.

Hon. Treasurer: Hon. Secretary:

DOROTHEA F. SANDERS, M.Sc. MARGARET I. R. SCOTT, M.Sc.

Hon. Librarian:
F. S. COLLIVER

Hon. Editors:

S. T. BLAKE, M.Sc.
GEORGE MACK, B.Sc.

Members of Council:

M. J. MACKERRAS, M.Sc., M.B., Professor A. L. REIMANN, D.Sc., Ph.D.,
J. H. SIMMONDS, M.B.E., M.Sc., Professor W. STEPHENSON, B.Sc., Ph.D.,
Professor L. J. IJ. TEAKLE, B.Sc.Agr., M.S., Ph.D.

Hon. Auditor:

L. P. HERDSMAN.

Trustees :

F. BENNETT, B.Sc., Professor W. H. BRYAN, M.C., D.Sc.,
E. O. MARKS, M.D., B.A., B.E.

CONTENTS

Vol. LXIII.

No. 1. — Form or Function. By M. F. Hickey. (Issued separately,

29th June, 1953)

No. 2. — Studies of the Life Histories of Some Queensland Blattidae

(Orthoptera). Part 1. The Domestic Species. By Pauline

Pope. (Issued separately, 6th July, 1953)

No. 3. — Studies of the Life Histories of Some Queensland Blattidae

(Orthoptera). Part 2. Some Native Species. By Pauline

Pope. (Issued separately, 6th July, 1953)

No. 4. — Parasites of the Bandicoot, Isoodon obesulus. By I. M. Mackerras,
M. J. Mackerras and D. F. Sandars. (Issued separately,

6th July, 1953)

No. 5. — A Study of Diphyllobotliriidae (Cestoda) from Australian Hosts.

By Dorothea F. Sandars. (Issued separately, 6tli July, 1953)

No. 6. — Two New Metastrongyle Lung-worms from Australian Marsupials.

By M. Josephine Mackerras and Dorothea F. Sandars. (Issued
separately, 6th July, 1953)

Report of Council

Abstract of Proceedings

Pages.

1-22

23-46

47-60

61-64

65-70

71-76

vii.

«»

PROCEEDINGS

OF THE

ROYAL SOCIETY

QUEENSLAND

FOR 1951

VOL. LXIII.

ISSUED 17th AUGUST, 1953

PRICE: TWENTY-FIVE SHILLINGS.

Printed for the Society
by

A. H. TUCKER, Government Printer, Brisbane.

NOTICE TO AUTHORS

1. Each paper should he accompanied by the author’s name, degrees and official
address.

2. Papers must be complete and in a form suitable for publication when com-
municated to the Society and should be as concise as possible.

3. Papers must be accompanied by a synopsis prepared according to the direction
given on the inside of the back cover.

4. Papers should be in double-spaced typescript on one side of the paper with
ample margins.

5. The use of italics in the text should be restricted to generic and specific names,
foreign words, and titles of periodicals.

The cost of author’s corrections to proof above what the Council considers a
reasonable amount, must be borne by the author.

7. Unless otherwise specified each author will be supplied with fifty separate
copies of his paper. Any number exceeding this may be obtained at approxi-
mately cost price.

8. All references should be listed at the end of each paper and arranged
alphabetically under authors’ names, e.g.,

Keilin, D. (1929). Proc. Roy. Soc. B., 104, 207.

Lesage, P. (1895). Ann.Sci. Nat. Bot., 1, 309.

The corresponding references in the text should be:

“ Keilin (1929) ”, “Lesage (1895) ”.

9. The size of the printed plate will not exceed 8 in. x 4^ in., and drawings may
be to this size, or preferably to a convenient small multiple thereof. The
effect of the necessary reduction on lettering and fine detail should be borne
in mind. Text figures should be drawn for reduction to a width not
exceeding 4 in.

Vol. LXIII. No. 1.

Proceedings of the Royal Society of
Queensland.

PRESIDENTIAL ADDRESS.

FORM OR FUNCTION.

M. F. Hickey.

[Delivered before the Royal Society of Queensland , 2nd April, 1951.)

Introduction.

It is a traditional practice in our Society that the retiring President
should close the annual meeting with an address or sermon — a task which
I find more difficult when I recall (as you undoubtedly will) the addresses
of previous Presidents, lingering memories of whose learning and rhetoric
may lead you to make odious comparisons.

As my principal work has been as a teacher of Anatomy and as I have
for many years been interested in the wider implications of biological and
general scientific problems, I hope that you will bear with me if I attempt
in this address a two-fold task ; to deal first with some problems inherent
in the relationship of anatomy and physiology as University subjects,
and secondly to discuss some of the more general problems, verging on or
including the philosophical, which arise from a consideration of the relation-
ship between form and function, in both biological and non -biological
fields.

Part I.— THE EDUCATIONAL PROBLEM.

The title of my address was chosen, partly because it is sufficiently
elastic to allow me some latitude in discussion, and partly because for some
time there has been in the biological studies in general a real divorce, and
in some cases an antagonism, between these two aspects of a single
problem Especially is this so in human anatomy and human physiology.

(a) The Correlation of Anatomy and Physiology.

Anatomy in general is concerned with structure, and physiology with
function. Time does not permit me to record the historical causes of the
divorce between these two disciplines ; however, the divorce of the two
interdependent aspects of a single subject is a relatively recent one, and
is responsible for a considerable wastage of time and effort and loss of student
interest in the medical course. It is also responsible for the fact that medical
students and others tend to keep their studies of anatomy and physiology
in “ water-tight ” compartments. However, the relative neglect of function,
both internal and external, is found in biology in general ; and I think
that the teachers of zoology and botany in our Universities might well ask
themselves how many of the thousands of men and women who have
passed through their hands still have a vital and abiding interest in
biological inquiry, even as an avocation.

2 PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

In most medical schools, and this applies particularly in Australia
(I don’t know what the position is in the veterinary schools, but have no
reason for thinking that the situation is any better there), there is little
or no correlation between anatomy and physiology. Into the present
reasons for this state of affairs, which are not flattering either to the
anatomist or to the physiologist, we cannot enter here; but I wish to say
a few words about its effects.

The separation of the two subjects is seen in many ways : the com-
petition for working time in the medical curriculum ; the lack of co-operation
between departments ; the tendency — nay, the necessity — for the student
to treat the two subjects as completely separate ones, and his failure con-
sequently to develop a “ total ” view of human structure and function ;
the real failure to evoke interest in and love for the subjects ; the complete
absorption of the student’s time in his strictly professional studies, with
a resulting loss of time, opportunity and inclination for wider education.

If we were to suggest to a mechanical engineer the complete separation
of the structural and functional aspects of his subject, he would think
we were joking ; yet it is a common practice in medical schools not only to
have anatomy and physiology taught completely independently from the
administrative point of view, but to tolerate a spiritual separation of the
departments, a far greater handicap than a merely geographical separation,
as is evidenced by the almost complete ignorance of each department
of what is being taught in the other, and a lack of synchronization of treat-
ment of material. So bad is this last feature, that the student may be
called upon to make a detailed study of function in relation to tissues,
organs and organ systems, before he has had the opportunity to acquire
the essential knowledge of structure. As a result of this, students find that
they are studying for two different examinations, and often studying
conflicting views on the same subjects. Any suggestion that this situation
should act as a stimulus to the critical powers of the student reveals a
complete failure to appreciate the realities of the situation. Assuming even
a very high standard of intellectual equipment in the student, this view
fails to take account of the fact that, under the present conditions, the
student has not the critical equipment either of factual knowledge or of
training to enable him to make a judgment between competing views.
It is realised of course that there are fields of anatomy of considerable
value in clinical work which do not demand excessive physiological treat-
ment, and fields of physiological study the anatomical basis of which is
relatively restricted. However, this cannot be considered an impediment
to a careful integration of the two fields of study which would involve
close co-operation between the departments involved and a continual
correlation of the subjects taught.

Evidences of a new attitude to the problem of reconciling the claims
of anatomy and physiology are not wanting, and, in this, the attitude is
important rather than the details of particular plans. It may interest
you to know that an attempt has been made to deal with the problem
in the University of Birmingham, where anatomy and physiology have
been integrated and a correlated course devised enabling the structure
and function of the body to be dealt with as one subject (Zuckerman,
1947).

As T. P. McMurray, Emeritus Professor of Orthopaedics in the
University of Liverpool, said (1949) : —

“ The idea is excellent ; through this arrangement the bald truths of
anatomy can be enlightened by the experimental work carried out in the
physiological section, and the dead structure of the dissecting rooms can be
seen to function in a living medium.”

FORM OR FUNCTION.

To devise and implement such a plan would, of course, make demands
on all the teachers involved for a study of their subject in relation to the
whole curriculum, in relation to other subjects, and in relation to the
student’s needs — rather a novelty in the building of a medical curriculum.
For several years, the working out of such a plan has been advocated in
our University, and the Faculty of Medicine has approved of the idea in
principle. But practically nothing has been done to cope with a situation
which, quite frankly, both in the fact of and in the reasons for its existence,
is a reproach to our University in this half of the twentieth century. In
the other Australian medical schools the situation appears to be quite
as bad.

Into the causes of this failure to attempt, let alone achieve, this fusion
there is no point in entering here. I would merely say that the fault appears
to lie equally with the anatomists and the physiologists, as evidenced in
the building of University curricula and in the strictures passed by each
group upon the other.

In the medical curriculum there is competition for time-table space
between the anatomists and the physiologists, and when the protagonist
of one or other of these disciplines fails to secure what he considers to be
adequate formal time-table space, he tends to seek, by large reading
assignments and the like, to secure it from the student’s “ free ” time.
In this competition, the anatomists have tended to be on the defensive
and the physiologists on the offensive. Medical teachers and students
are from time to time informed by physiologists that “ anatomy is relatively
unimportant in the medical course, and therefore in medical science and
practice.” “Anatomy can be taught in a few easy lessons.” “ Morphology
is finished as a research subject.”

We may ask ourselves whether there is any truth in these statements,
and in their implied corollaries : that physiology is the only subject of
importance in medical science and practice ; that physiology needs and
deserves much more time than anatomy ; that physiology is the only
worthwhile field of research. The claim that morphology is finished as a
field of research has two aspects ; one, explicit, that the whole field of
morphological investigation has been worked out, a claim that is difficult
to reconcile with the great deal of work still being done both in microscopic
and gross anatomy, even if we confine our attention to the recent important
large scale re-investigations of major problems in surgical anatomy ; the
other aspect, implicit, is that the availability of research fields is the most
important criterion of the importance of a subject in a medical school,
or in a University in general.

This latter notion arises from a misconception of the function of the
University teacher and of the functions of a University. Although it is
true that the ideal University might be described as a training ground for
critical thought, the fact is that this ideal function is an accident of the
type of studies undertaken at Universities in the first few centuries of then-
existence. The Universities began as professional training schools : rightly
or wrongly this is also one of their principal characters to-day. From
the unfortunate over-emphasis on research as a competitor with, rather
than as a stimulus to, teaching stems the fact that appointment to and
advancement in University posts goes too much by a reckoning of paper
output rather than by educational productivity. There is also the fact
that University teachers too often forget that the student must be trained
in thinking and in scientific method, and must he helped to enlarge his
field of knowledge. This requires time and thought. We should remember

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

that there is an advancing edge to the student’s knowledge as well as to
that of the research worker ; and in general, both academically and socially,
the former is the more important.

Turning to the question of the importance of anatomy, of structure,
in the medical course. Assuming that the separation of structural and
functional aspects is valid, the suggestion that anatomy is of minor
importance in the medical course and therefore, one must suppose, in
general practice, is simply a blind perversion of the facts.

It is an inescapable truth that there is no other basis for the study of
function and of disturbed function, than a thorough knowledge of relevant
structure. As Professor McMurray put it recently (1949) :

“ The student cannot possibly know too much of that subject (Anatomy) :
It is just as important for the Doctor to know every part of the human organism
as it is for the Engineer to know every nut and bolt of the machine for the repair
and maintenance of which he is responsible. The only question then is whether
the time spent in anatomical teaching could be rearranged to equip the medical
student more fully for his life-work.”

A sound knowledge of normal living anatomy is the fundamental
basis of the examination of a patient and is essential for a great deal of
treatment. There should be no need to labour this point any further ;
but relevant questions are, what should be the scope of anatomy, what
facts or groups of facts can be omitted from the course, what others could
with advantage, be stressed or elaborated ? These questions, of course,
might appear meaningless to the pure anatomist to whose mind the subject
of anatomy consists of a mass of facts each of which is of equal value. Let
us recognise clearly, however, that in medical schools at least, we are training
for the practice of medicine and not for a life-long pursuit of anatomy.
Into these problems of selection and emphasis of material this is not the
occasion, nor is there the time, to enter ; but I should like once again to
emphasise the view that so far as the medical student is concerned, one
great problem facing the teacher of Anatomy, qua teacher, is the problem
of method.

(b) Teaching Methods in Anatomy.

I am now going to discuss some problems of technique in anatomical
and general morphological instruction. Are there relatively easy and
intellectually satisfying methods of teaching structure, and have these
been sufficiently exploited ? I think that the answer to the first question
is yes, and the answer to the second is a very definite no.

This brings me, then, to the question of teaching technique, mainly
in human anatomy. I am not at the moment concerned with how much
detailed anatomy should be taught to medical students, for that is not of
importance in this discussion, but rather with the consideration of whether
present methods of instruction are adequate. . Here we may view the
matter historically to see how the present situation has arisen. As you
know, Physiology and Biochemistry are relatively recent in their full
separate development ; and traditionally medical students were concerned
with two main activities — dissection of the cadaver, and “ walking the
wards.” A relatively enormous amount of time was allotted to dissection,
the body being dissected in detail twice. The student was given a dis-
secting manual and a cadaver, and set to work on his own voyage of
discovery. To a great extent this method still persists ; but even if the
time at the student’s disposal were unlimited, I should still consider this
approach to the subject a very poor one.

FORM OR FUNCTION.

Again, as J. D. Bernal (1939, pp. 78-9) has remarked, University curricula
have grown by a process of accretion and compression, with very little
pruning or attempt at correlation. In the last half century, a whole new
field of work in physiology and biochemistry has developed, making more
demands on the time and attention of students, but with little accompanying
alteration in the scope or method of anatomy.

Are we able to make the approach to dissection one which would
produce a much better informed dissector ? The answer, I think, is yes.
The first and most important thing to do is, both by ordinary didactic
lectures and by demonstration of specimens and of models, to impart
to the student a thoroughly sound knowledge of the general architectural
problem. In most cases, this involves a complete reversal of the ordinary
dissecting room procedure. Instead of working from the skin inwards,
we should work from the bones outward, thus preparing the bony skeletal
framework in which we are to place muscles, glands, nerves, and vessels.
This involves a considerable degree of instruction antecedent to dissection ;
but it also means that when the student proceeds to his dissection he does
so reasonably well-informed on the general build of the part he approaches.
This will ensure that his dissection will be a much better one, much more
rapid, and in effect a more detailed revision of the preliminary instruction.
It is during this second stage that such details as are considered necessary
can be imparted to the student.

It may be objected that by this method too much is done for the
student. We could admit the validity of this criticism, if we also admitted
that it is not desirable for the student to be a well-informed dissector.
What is aimed at in the method suggested is a rapid acquisition of a sound
basic knowledge of a region, so that the student may make his dissection
of the region a much more profitable one. It has also the great advantage,
especially in studying joint mechanisms, one of the most neglected fields
in the anatomical training of medical students, of allowing the functional
aspect to be dealt with early, thus giving more point to the study of joint
and muscle structure.

What accessory devices have we to assist the teaching of structure ?
The following may be enumerated — a series of dissections showing the
principal masses of a region arranged on a basis of bony structure from
within outwards, correlated with the systematic study of cross-sections ;
this to be supplemented by special dissections of parts where the details
are considered to be of importance in medical practice ; models which can
be of enormous assistance in gross anatomy, embryology and histology ;
and carefully selected sets of diagrams, x-rays and films.

The use of the above method of approach and of the visual aids
suggested, will mean that a student can be put very rapidly in possession
of the significant facts of structure as a prelude to or as an accompaniment
of the study of function. The result would be considerable saving in time
and mental energy. It would ensure a much better dissection and a much
sounder knowledge of structure, it would allow greater time for revision,
and it would allow a much more knowledgeable approach to the study of
normal and disturbed function.

Of course, it may be claimed that technique is easy in theory but
difficult in practice ; so I now wish to mention some simple teaching devices,
a wider application of which would make the acquisition of knowledge of
structure easy and interesting.

6 PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

[The following part of the lecture was illustrated with lantern slides and
films.]

(1) A method for practical histology which aims to help the student
to become well-informed on general build before he approaches the problem
in detail. One of the great difficulties in large practical classes in micro-
scopic anatomy, especially with junior students, is to ensure that all students
have a reasonably clear idea of the appearance of structures for which
they are searching. Coloured transparencies of successively greater magni-
fications photographed directly from class slides can be used to show first
the whole mount of tissue , so that the general arrangement can be explained
and certain areas pointed out for study ; then a low power view of a fairly
large segment of a special structure ; and in later slides high power views
can be shown in which considerable detail can be made out and demon-
strated to the student. Other parts of this and other material can be
treated in the same way. This method, if properly applied, will make
the time needed for the study of pure structure very much less ; but it
will ensure that the whole class has the opportunity to see what the structure
is, and provide more time to make a thorough study of details and so
achieve a higher degree of proficiency in the subject. In any rational^
planned medical course this will have three advantages : in the first place,
it will give a surer structural basis for the study of function ; it will give
more time for the study of function ; and it will provide a sound basis
for work in histo-pathology.

(2) A device by no means new, but poorly exploited, is the use of
simple models. Home-made models, constructed without great difficulty
and in a short time, give the student an incomparably better idea of the
general build of the organism or part than do the usual drawings in the
textbook, for the three-dimensional qualities can be thoroughly appreciated.
Lantern slides made by photographing such models are generally superior
to those made by photographing a drawing of a model. Of course, this
method applies particularly well in embryology, but it is also very useful
in gross anatomy, for by means of a series of simple models, the fundamental
architecture of a region can be demonstrated with ease.

[Some home-made models were displayed to give an idea of the usefulness
of the material, especially in the teaching of Embryology.]

(3) The third device is the carefully planned application of a well-
known technique, the use of ordinary lantern slides in gross anatomy
instruction. As a preliminary to the study of a part, the general relation-
ships can be discussed with a large class by means of a careful correlation
of cross sections and dissections in flat planes. This can be supplemented
by the study of models and of prepared dissections. It is essential that the
slides be very clear and not too numerous.

(4) One could continue the enumeration and illustration of these
and similar devices, but I will conclude this part of my talk with a few
remarks on a technical device of great value in the teaching of living anatcmy,
the film, silent or with a sound track.

Again, it must be remembered that the film is only a means to the
understanding of the actual working body ; that there is the danger of
the over-use of films, especially excessively long ones ; there is the danger,
too, of the student playing a purely passive part in the process. These
dangers and others are known and can be coped with. Here I wish to say
something about the value of films in anatomical teaching.

FORM OR FUNCTION.

The films may be roughly divided into four classes :

{a) Strip films showing successive stages in a dissection, either as a
prelude to, or as a revision of, actual dissection.

(6) Films of the animated cartoon type showing embryological develop-
ment and function.

(c) Ordinary films of the living body showing the movement of joints
and permitting the analytical study of joint movement, &c.

(d) Films showing successive stages in joint movement or in the passage
of material through tubular structures as visualised by x-rays.

(c) and (d) are types which are easily prepared in an Anatomy depart-
ment equipped with an x-ray apparatus and suitable photographic equip-
ment. I will first show you a film showing the passage of material through
a tubular structure : a barium meal passing through the stomach. Then
will follow three films to illustrate joint movement in ankle, elbow and
forearm.

[Four short films were shown.]

It may interest you to know how such films can be prepared. There
are two methods. One is to photograph directly the movements af
visualised on the fluoroscopic screen. Technically this is somewhat difficult,
but necessary for the study of the movements of tubular structures. The
other method, of especial use in the study of joint movement, is to take a
series of x-ray pictures in a considerable number of consecutive positions,
then photograph these on to a continuous film by using the appropriate
number of exposures for each successive position so as to give the illusion
of practically continuous movement.

[Illustrated with a series of five lantern slides made from x-ray pictures
of five consecutive positions of shoulder joint movement. By increasing con-
siderably the number of intermediate positions there is provided the raw material
for a short film, which can be run as an unbroken ring, allowing a complete
visualisation of the movement and, in successive stills, a means of analysis of
relative angular movement of joint components.]

The advantages of this method are too obvious to need stressing.
But here again I cannot too strongly emphasise that this method is only
a means to an end — the understanding of the function of the joint in the
living body.

I would venture to suggest that somewhat similar devices would be
of enormous value in the teaching of physiology and, with due respect, I
would also suggest that the carefully prepared film showing some of the
routine animal experiments (if indeed these must be used) as a preparation
for the actual work by the class would be a very great assistance both to the
students and to their instructors. It is a far from uncommon experience to
find that the experimental result is nil, due to the imperfect technique of the
untrained and ignorant student.

It will no doubt occur to you that much the same problems of emphasis
and method will arise in biology in general. However, both the potential
dangers and advantages are greater here, for owing to the fact that both
aspects are usually taught in the one department, it may be possible to
find biology courses in which there is a careful correlation of structure
and function. Nevertheless the very real danger exists that there will be
an exclusive emphasis on morphology, with little or no treatment of
physiology, ecology and the like, as so wittily and trenchantly stated by
William Morton Wheeler (1923); and there is the danger that an excessive
preoccupation with physiology proper or with genetics may deprive the
student of the opportunity to get to know the animals themselves.

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

I well remember how after I had graduated in Medicine, I was given
a copy of Wigglesworth’s small book on “ Insect Physiology ” by a
friend, and how, after a casual glance at the first few pages, I sat down
enthralled to read the book through completely. As I went on with the
reading, the question kept occurring : why were we not given some similar
approach to our study of insects and other animals in our first year of
biology ? Here was a key to the understanding of structure in terms of
function. To what extent function is now being correlated with structure
in University courses in biology, I do not know at first hand, but I am
inclined to doubt whether nearly enough time and attention is given to
the functional illumination of structure as is needed to secure full interest
and understanding on the part of the student.

I know, of course, that a certain amount of ecology is now done ; a
small amount, owing to the difficulties of time-tables. This is important,
but is not sufficient in itself ; nor does there seem to be enough time and
attention devoted to the general functional aspects of the animal types
which are studied in the practical classes. This is particularly true of
locomotion, a subject easy to study, if not on the living animal, then in
films. I feel that this is only partly, and to a minor degree, due to a lack
of appreciation of the importance of function, and I am not suggesting
that the study of structure be sacrificed to the study of function. Far
from it, for we would not wish to have the reproach levelled against
zoologists excessively preoccupied with minute physiological and bio-
chemical problems, that the study of aniimls is unfashionable among
them.

It will be apparent that in dealing with the internal correlation of
the biological sciences and with the technical problems of teaching in them,
we can apply the same principles and have recourse to the same aids as
in the teaching of anatomy, and expect to obtain the same advantages.
I do not think that there is any need to labour the point.

Probably the divorce of structure and function has never been as
marked in general biology as in the medical subjects, unless perhaps in
comparative anatomy. Here, however, there are signs of a widening of
the scope of the subject/, as is evidenced in two outstanding text-books
recently published. In his new work on comparative anatomy, Professor
Romer (1949) has paid considerable attention to the cognate sciences,,
including physiology, as well as to essential anatomical or morphological
aspects, producing a work which, according to William L. Straus (1950)
may play a considerable role in the rehabilitation of this branch of biology.
If comparative anatomy is to recover lost ground and prosper, it must
become broader in scope ; in fact it must become comparative biology,
instead of the usual narrow comparative anatomy or morphology ; and
Romer’s book seems to have the necessary qualities.

This approach seems to be even more fully used by Professor J. Z.
Young in his recent book (1950) which shows a much needed “ holistic ”
approach to the problem, with structure and function treated together.
In fact, his method of treatment “ challenges the appropriateness of the
conventional distinction between the two subjects.” He deals with the
whole working organism, and provides not simply a functional morphology
nor a physiology with an anatomical substratum. Professor Young’s
expressed intention is to treat of the fife of vertebrate animals in its most
complete synoptic sense, its origin and secular changes of complexity,
the means by which animals cope with the hazards and contingencies of

FORM OR FUNCTION.

living and staying alive, the varieties of truce that accommodate the chronic
enmity of the environment, and everything else that is entailed by the act
of living. “ The central fact of biology ” says the author, “ is that life
goes on.”

(d) Some General Considerations.

If the example of the University of Birmingham in anatomy and
physiology is more widely tried out, and if the attitude of Romer and
Young becomes more generally adopted, we may have some hope that
the leaven will spread rapidly amongst University teachers and have some
influence on the anatomists and the physiologists, so that both may come
to realise that they are guiding students, and not teaching subjects. It
may even come about that medical faculties and their committees charged
with the construction of curricula, may give some considerable degree of
attention to this fundamental problem of correlation. We realise* of course,
that this may make serious demands on the egoism of both anatomists
and physiologists. It may also mean that each will have in some way or
other to keep themselves ‘ au fait ’ with what is going on in each other’s
fields of work. It may mean even more that some serious attention will be
given to the scientific approach to the problems of curriculum structure
and of teaching.

I make no apology for having spoken at some length on the problems
of correlation of subjects and teaching techniques. We have too long
had an excessive tenderness for professors and lecturers whose main
qualifications have been the passing of examinations and the writing of
papers. Let me make it clear that I am not denying the importance of
research work, but I do consider that we should see it in its proper per-
spective. Whether we like it or not, in medicine, dentistry, veterinary
science and applied biology, we are concerned with training students for
professional careers in which a working knowledge of the subject is of
great importance for the performance of everyday professional tasks.
This preparation is both historically and sociologically a University work.

It may be claimed, and often it is, that if we set about devising curricula
on a planned system, and if we look for newer and easier methods by which
the student may master the material essential for his professional work,
we are “ spoon-feeding ” him, that we are getting away from the true
spirit of the University, and that we will destroy his initiative. Is there
any substance in this criticism ? Frankly, I think not.

If we are able to devise better methods of instruction, especially on
the basic principles of a subject, we can make much heavier demands on
the student in terms of performance. Let us look at it in this way. If
we are instructing students in anatomy, including histology, as a preparation
for their study of physiology, then surely in all common sense we should
teach as much as is possible of the common ground of structure and function
in the one course. Further, the devising of techniques of instruction,
which will make as easy as possible the acquisition of a thoroughly sound
knowledge of structure, will not only make the student more interested in
his subject, due to his more rapid appreciation of it, but also will give him
much better intellectual equipment and much more freedom of time and
mental energy for the study of function, which many physiologists will tell
us is of overwhelming importance. It may give the student some free time
in which he can be encouraged to study the history and structure of the
society in which he is to do his life’s work, to become acquainted with some
at least of the literary and artistic life of the world.

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

We must not lose sight of the fact that all of these devices are merely
a means to an end, to a knowledge of the build and working of the living
body ; but the means are of great importance not only in this particular
field, but in the wider fields of education.

A consideration of methods is only part of a scientific approach to the
problems of teaching, and teaching is teaching whether in a University
or in a secondary school. It has problems of technique as well as content
and correlation. The importance of a high standard of skill in University
teachers as such is now being stressed in widely diverse quarters. For
example the former Vice-Chancellor of the University of Melbourne, Sir
John Medley (1950) leaves no doubt about his views : —

“ I feel sure that of late years we — and I speak now of University theorists
in general — have tended unduly to exalt research at the expense of teaching,
and that the true picture of a University’s function has been blurred thereby. . .
And yet to hear some of the extremists talk you would come to the conclusion
that teaching was nothing but an unfortunate but inevitable excrescence upon
the body academic, and that nobody should be appointed to a University staff

unless his research record was of first rate calibre The teacher

is the backbone of any true University, and there will always be teachers of
first rate capacity whose talents do not lie in the direction of original work and
will be sadly wasted if pressure of academic opinion forces them to devote time
to so-called research that could be better spent in their proper job.”

Do not imagine, however, that Sir John regards research as of no
account ; far from it, as a perusal of his whole address will make clear.

Now Sir John Medley is an administrator, and perhaps you may think
that his views are prejudiced in favour of administration and teaching.
Let me quote to you the words of one of the leading researchers of our
time, Sir Macfarlane Burnet (1948), who, in discussing the relation between
teaching and research in the University, writes as follows : —

“ There is a widely current attitude that the only activity of a professor
which matters is his research work : teaching and administration are time-
wasting obstacles which are necessary but unfortunate concomitants of the
appointment.”

“ This seems to be a highly unhealthy condition, which, like most things
human, does not work out quite so badly as it ought to. It is a rather naive
and socially untenable attitude to assume that part-time research, which,
because it is part-time, must in most instances be of relatively unimportant
character, is to be regarded as of greater importance than effective teaching.
Good education, with which must be included proper selection of those fitted
to benefit by it, is the most important requirement for the production of medical
men and scientists. The effective organization of a University department,
so that it can fulfil all its functions of teaching, research and outside advice,
is a full-time job that is more important than any but the rarest of advances
in research.”

“ This may seem a highly heretical statement worthy only of those who
regard a university simply as a school for higher vocational training ; but as
one who can hardly be accused of ignorance about the values of fundamental
research, I feel strongly that it should be seriously considered by anyone con-
cerned with attempts to remedy the present difficulties of Australian
universities.”

Professor Burnet then outlines the ideal structure of a University
department in one of the Sciences.

“ The head of the department would have final responsibility for all its
activities but would spend no significant proportion of his time on personal
bench investigations. The professor’s primary interest would be in effective
teaching, using this in the broadest sense, and in the organisation of his depart-
ment to provide this more and more efficiently. His activities at the research
level would be essentially research into teaching methods, the devising of visual
aids, improvement in laboratory training, the assessing of capacity for specialised
work in students : in other words, educational research applied to the teaching

of his science.”

I should like to quote this article at greater length, but time permits
no greater quotation than I have so far made.

FORM OR FUNCTION.

Part II.— THE PHILOSOPHICAL AND ETHICAL PROBLEMS.

(a) In Biology.

With your indulgence I now propose to deal briefly with some general
problems that arise in biology concerning the meaning of the relation of
living function to living structure and the relation of both of these to a
wider scheme of things. For example, it is held by some, and denied by
others, that some aspects of modern biology have been destructive of
•ethics.

It has been suggested, for example, that the insistence on the struggle
for survival and on the survival of the fittest in nature has made it difficult
for us to believe in the possibility of finding a worthwhile ethical system ;
and that this is due to the work of men such as T. H. Huxley. This belief
has arisen from a misconception or a plain misrepresentation of 19th century
biology.

One of those who declare, and presumably believe, that Tennyson’s
phrase of “ Nature red in tooth and claw ” was adopted by tin immediate
exponents of Darwinism, is that distinguished anatomist, Professor Wood
Jones ; and in a recent book (Wood Jones, 1942) he sets out to propound
that biology everywhere shows evidence of design and of purpose ; and
incidentally, that the struggle for survival is net the keynote of the world
of nature, but that mutual aid and mutual support are the fundamental
characteristics of nature.

It is not possible here to deal in detail with this thesis, but I think
it is proper that certain aspects should be mentioned. Wood Jones rightly
points out that Darwin realised there was mutual dependence, as well as
competition, everywhere in nature. However, he goes on to say (p. 45) : —

“ The pictures of relentless competition, of bloody strife, and ultimate
elimination of the weak in favour of the strong made great appeal to Huxley
in England and to Haeckel in Germany, and by them they were handed on to a
receptive public incapable of estimating their true value.”

I think that Wood Jones is right in stressing the implication of
dependence of living things upon each other, as did Kropotkin (1902,
1919) ; but I think he is in error in declaring simply that “ nature is a
series of harmonies in which living things play mutually dependent parts,”
and that increasing population does not produce inevitably and funda-
mentally a competition for survival, the struggle for existence, both within
groups and between groups. He must be blind to what is going on in the
world about him, both human and non-human.

However, I am not concerned here to support this theory of the struggle
for existence in Nature, but to draw attention to and deplore Wood Jones’
implicit misinterpretation of T. H. Huxley ; for he would leave us with
the idea that Huxley believed and taught that the struggle for survival
should be the keynote of the human situation, that it was the teaching
of Huxley and his school which was responsible in great part for the present
chaos in theological and ethical beliefs.

The fact of the matter is that Darwin never suggested that we should
transfer the operations of the rest of the animal kingdom to the field of
human conduct ; and indeed it was T. H. Huxley (1895) who emphasised

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

most strongly that the cosmic process in nature must be subdued to the
ethical process. Let me quote from Huxley himself (pp. 80-83) : —

“ There is another fallacy which appears to me to pervade the so-called
‘ ethics of evolution.’ It is the notion that because, on the whole, animals and
plants have advanced in perfection of organisation by means of the struggle
for existence and the consequent ‘ survival of the fittest ’ ; therefore men in
society, men as ethical beings, must look to the same process to help them towards
perfection. I suspect that this fallacy has arisen out of the unfortunate ambiguity
of the phrase ‘ survival of the fittest.’ ‘ Fittest ’ has a connotation of ‘ best ’ ;
and about ‘ best ’ there hangs a moral flavour. In cosmic nature, however,
what is ‘ fittest ’ depends upon the conditions ”

“ Men in Society are undoubtedly subject to the cosmic process. As among
other animals, multiplication goes on without cessation, and involves severe
competition for the means of support. The struggle for existence tends to
eliminate those less fitted to adapt themselves to the circumstances of their
existence. The strongest, the most self-assertive, tend to tread down the weaker.
But the influence of the cosmic process on the evolution of society is the greater
the more rudimentary its civilisation. Social progress means a checking of the
cosmic process at every step and the substitution for it of another, which may
be called the ethical process ; the end of which is not the survival of those who
may happen to be the fittest, in respect of the whole of the conditions which
obtain, but of those who are ethically the best.”

“ As I have already urged, the practice of that which is ethically best- —
what we call goodness or virtue — involves a course of conduct which, in all
respects, is opposed to that which leads to success in the cosmic struggle for
existence. In place of ruthless self-assertion it demands self-restraint ; in
place of thrusting aside, or treading down all competitors, it requires that the
individual shall not merely respect, but shall help his fellows ; its influence is
directed, not so much to the survival of the fittest, as to the fitting of as many
as possible to survive. It repudiates the gladiatorial theory of existence. It
demands that each man who enters into the enjoyment of the advantages of a
polity shall be mindful of his debt to those who have laboriously constructed
it ; and shall take heed that no act of his weakens the fabric in which he has
been permitted to live. Laws and moral precepts are directed to the end of
curbing the cosmic process and reminding the individual of his duty to the
community, to the protection and influence of which he owes, if not existence
itself, at least the life of something better than a brutal savage.”

“It is from neglect of these plain considerations that the fanatical
individualism of our time attempts to apply the analogy of cosmic nature to
society ”

“ Let us understand, once for all, that the ethical progress of society
depends, not on imitating the cosmic process, still less in running away from it,
but in combating it.”

One would think that there could scarcely be a plainer statement
of the contrast between the ethical and cosmic processes ; you have your
choice of reading Huxley’s ideas in his own words or Wood Jones’ pre-
judiced mis-statement of those ideas.

However, Huxley realised quite well that human affairs show a struggle
between our cosmic and our ethical nature, one that will be as long lasting
as human life, but one in which will and intelligence will help the ethical
process to curb the cosmic, although with difficulty (p. 85) : —

“ Moreover, the cosmic nature born with us, and to a large extent necessary
for our maintenance, is the outcome of millions of years of severe training,
and it would be folly to imagine that a few centuries will suffice to subdue its
masterfulness to purely ethical ends. Ethical nature may count upon having
to reckon with a tenacious and powerful enemy as long as the world lasts. But
on the other hand, I see no limit to the extent to which intelligence and will,
guided by sound principles of investigation, and organised in common effort,
may modify the conditions of existence, for a period longer than that now
covered by history. And much may be done to change the nature of man himself.
The intelligence which has converted the brother of the wolf into the faithful
guardian of the flock ought to be able to do something towards curbing the
instincts of savagery in civilised men.”

FORM OR FUNCTION.

If I have quoted rather fully from Huxley, it is because it seems
necessary to make clear once again what Huxley did say, especially in
these times when too many prominent publicists, theological and other-
wise, blame the moral ills of the world on “ science ” or “ Darwinism,”
or “ scientific materialism,” in many cases without knowing what the
scientists concerned have said, and in some cases quite obviously mis-
representing their views.

It should be clear from the paragraphs quoted that Huxley’s view of
the ethical process as supplanting the cosmic process has implicit in it
the notion of interdependence ; indeed the ethical process can develop
only with an increasing recognition of this interdependence. Huxley’s
own words are explicit — “ Let us understand once for all that the ethical
progress of Society depends, not on imitating the cosmic process, still less
in running away from it, but in combating it.”

Another charge against 19th and 20th century biology is that it has
destroyed the “ comfortable ” assurance of an all-pervading design and
purpose in nature, with deleterious effects on ethics. Into the question
of whether there is purposiveness in nature, in evolution, and in human
life, we cannot enter at length here. There is no doubt about the views
of some of the leading biologists to-day on this matter. Alfred Homer
(1949), a leading American comparative anatomist and palaeontologist,
sums up modem views when he says that there is no evidence of design
or direction in evolutionary changes. Rather, the changes are due to
mutations ; the process of mutation seems to be merely one of blind random
change ; but, he says, “ Vertebrate evolution certainly appears to be a
process which has resulted in changes both useful and adaptive. How
can such results have come out of the mutation process ” ? In short, he
summarises modern views thus (p. 14) : —

“ Both observation and experiment indicate that even small mutations
in an adaptive direction have a distinct survival value, and may become dominant
in a species in a short time. This natural selection of such a random series of
mutations as have adaptive value would appear to be a major mechanism of
evolutionary change.”

The question is discussed by Julian Huxley in his “ Evolution : the
Modern Synthesis ” (1942), and so well discussed (pp. 576-8) that I will
quote rather fully :

“ The ordinary man, or at least the ordinary poet, philosopher, and
theologian, is always asking himself what is the purpose of human life, and is
anxious to discover some extraneous purpose to which he and humanity may
conform. Some find such a purpose exhibited directly in revealed religion ;
others think that they can uncover it from the facts of nature. One of the
commonest methods of this form of natural religion is to point to evolution as
manifesting such a purpose. The history of life, it is asserted, manifests guidance
on the part of some external power ; and the usual deduction is that we can
safely trust that same power for further guidance in the future.”

“ I believe this reasoning to be wholly false. The purpose manifested in
evolution, whether in adaptation, specialization, or biological progress, is only
an apparent purpose. It is just as much a product of blind forces as is the
falling of a stone to earth or the ebb and flow of the tides. It is we who have
read purpose into evolution, as earlier men projected will and emotion into
inorganic phenomena like storm or earthquake. If we wish to work towards a
purpose for the future of man, we must formulate that purpose ourselves.
Purposes in life are made, not found ”

“Human purpose and the progress based upon it must accordingly be
formulated in terms of human values ; but it must also take account of human
needs and limitations, whether these be of a biological order, such as our dietary
requirements or our mode of reproduction, or of a human order, such as our
intellectual limitations or our inevitable subjection to emotional conflict ”

14 PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

“ But let us not forget that it is possible for progress to be achieved. After
the disillusionment of the early twentieth century it has become as fashionable
to deny the existence of progress and to brand the idea of it as a human illusion,
as it was fashionable in the optimism of the nineteenth century to proclaim
not only its existence but its inevitability. The truth is between the two
extremes. Progress is a major fact of past evolution ; but it is limited to a few
selected stocks. It may continue in the future but it is not inevitable ; man,
by now become the trustee of evolution, must work and plan if he is to achieve
future progress for himself and so for life.”

“This limited and contingent progress is very different from the deus ex
machina of nineteenth-century thought, and our optimism may well be tempered
by reflection on the difficulties to be overcome. None the less, the demon-
stration of the existence of a general trend which can legitimately be called
progress, and the definition of its limitations, will remain as a fundamental
contribution of evolutionary biology to human thought.”

One of the great problems of the relations of structure and function
is that involved in the elucidation of the methods of evolution, that is the
relation of the varied modifications which structures have undergone
and the accompanying functional changes. As Romer (1949) points out,
44 Most structural and functional changes in the vertebrate body are quite
clearly adaptive modifications to a variety of environments and modes of
life.” This would be accepted both by neo- Darwinians and their opponents
— omitting, of course, those who deny an evolutionary process. But how
these adaptations have been brought about is a matter on which there
have been, and are, several theories. These theories are briefly : —

(1) The naive suggestion of adaptation changes occurring as if the
animal “ willed ” them, or as if its needs or desires in themselves brought
new structures or structural changes into being.

(2) These theories are not far removed from some “ philosophical ”
theories of evolution which have had, and still have a certain vogue..
Theories of orthogenesis assume that there is some mysterious 44 inner
urge ” or 44 inherent tendency ” within the organism which brings about
these changes ; then there is the theory that evolutionary changes are
the result of the 44 design ” of some supernatural force. Since such theories
are non- scientific, they cannot be scientifically disproved ; but we are
at liberty to look for more reasonable explanations of evolution based on
known facts.

(3) A more plausible attempt at interpretation of structural evolutionary
changes was that first advocated over a century ago by Lamarck — a belief
that characters acquired through the effects of the environment or of use
and disuse could be transmitted to subsequent generations. As if one
might say : 44 If the giraffes’ ancestors stretched their necks for foliage
on high branches, the effects of this stretching would be transmitted to
their offspring, generation after generation, and an elongate neck gradually
developed in the hereditary pattern.” Or again : 44 If the snakes’ lizard
ancestors ceased to use their legs in locomotion, the cumulative result of
disuse would be that of eventual loss of the limbs.” This attractive theory
seems simple, reasonable and natural ; but its present standing is very
poor indeed.

(4) There is finally the neo-Darwinian view that the major mechanism
of evolutionary change is the operation of the natural selection of such a
random series of mutations as have adaptive value.

However, this last theory is not accepted universally. You are probably
well aware that in Soviet Russia it is laid down politically as well as
scientifically, that the environment is the principal modifying factor in
evolutionary changes. Into the merits of this belief it is impossible to

FORM OR FUNCTION.

enter here, but we are entitled to say that the well-substantiated persecution
of Mendelian geneticists in Russia has raised the controversy far above
the level of a scientific dispute, and I think we are also entitled to ask
what is the motive behind the crisis in Soviet biology. It has been suggested
by Darlington (1947) that a government which relied on the absence of
inborn class and race differentials in man as the basis of its political theory
was naturally unhappy about a science of genetics which relies on the
presence of such differences amongst plants and animals as the basis of
evolution and of crop and stock improvement : it was desirable to have
a theory of genetics interpreted and controlled by Moscow.

But the advocacy of Lamarckism is not confined to Lysenko and
his political backers. Fairly recently Wood Jones in “ Habit and Heritage 9r
(1943) sets out to re-establish Lamarckism, or, the transmissibility of
somatic modifications or acquired characters, i.e. “ features developed during
the life of the individual possessing it in response to the action of use or
environment.” Into the validity of Wood Jones’ thesis it is not possible
or necessary to go at this juncture. There are however, two points on
which I should like to comment. The first is his attitude to modern genetics
as is shown in the conclusion of his book, where derisively but with felicitous
style he likens the geneticist to the Chinese cashier manipulating his abacus.
He says (p. 99) :

“ I have, at times, been tempted to wonder if my unbounded admiration
for the facility with which the modern geneticist solves his problems, by manipu-
lating hypothetical little balls — called genes — along the bars of the chromosomes,
may be in part due to my astonishment at his dexterity and to my mystification
concerning the niceties of the working of his instrument. We have now a whole
race of scientists trained in the refined use of the abacus of heredity. Their
manipulation of the little balls of destiny has become an occupation demanding
extraordinary skill and a deal of specialized knowledge and literature. Beyond
any doubt they possess an instrument and a technique that enables them to
keep tally of the petty cash transactions of heredity. It is doubtless a mere
impertinence on the part of one who is not a professed, nor even an initiated,
geneticist to wonder at times if the great reliance placed upon the workings
of the abacus of heredity is not perhaps repressing the development of a possible
higher mathematics of inheritance. For undoubtedly there is a higher sphere
of inheritance and it must be sought, not through a microscope, not even by
the most nicely adjusted experiments, carried on for a short time in the short
life of one human being, but by a survey, incomplete though it must necessarily
be, of what Nature, with time unlimited at its disposal, has effected among
living things.”

“We may be running the risk of becoming blind to the whole range of the
possibilities of inheritance, if we concentrate too exclusively on the minutiae
of the means and modes by which in certain cases, it appears to be effected.”

Those of us who know something of the work of the modern geneticist
can assess this passage at its real value. We see in it many of the devices
of the advocate, not of the disinterested seeker after truth. We observe,
for example, the false modesty (“ my unbounded admiration ” ; “a mere
impertinence on (my) part ”) ; the pretence that here is a field of work
into which the writer is not qualified to enter (“ my astonishment ” ; “ my
mystification,” etc.) ; the mild derogation (“ petty cash transactions ”) ;
the implied untruth that geneticists are unaware of the necessity of studying
the “ higher sphere of inheritance ” because they “ concentrate too
exclusively on the minutiae,” etc. One could go on to point out the falseness
of the analogy of the geneticist with the abacus worker, the fact that many
first rate geneticists are also very keen naturalists and general biologists,
that the study of variation and genetics goes hand in hand with the study
of the causation of variation. But to do so would be a little pointless ;
we need not expand this analysis, for the explanation of Wood Jones’

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

views is to be found not in what he wishes to establish as scientific truth
but in what is his purpose in writing the small book from which the quoted
paragraphs are taken.

Wood Jones would have us believe that he prefers to be uninformed
of the whole field of modern morphological and physiological work in
genetics, so that he may cling to certain beliefs which he hopes are true.
He seems to have allowed his scientific attitude to be clouded over with
an adherence to certain philosophical principles, for in his first chapter
he suggests that what he calls “ the breakdown of ethical standards ”
is due in some degree to the widespread acceptance of the Darwinian
“ struggle for existence,” and rejection of the idea that acquired
characteristics can be transmitted, for he writes (p. 11):

“ Has not science — even in the most considered and orthodox pronounce-
ments of its most distinguished exponents — possibly led humanity astray in
the matter of responsibility in heredity ? There was a time when people believed
that the sins of the fathers were visited on the children. This belief was incor-
porated into the teaching that the parents need have some care of their ways
of life, lest their acquired shortcomings and moral lapses might be perpetuated
in their descendants. It led also to a general belief in the possible ultimate
betterment of mankind by a raising of the standard of parental mode of life
and well-being. Such hopes have been dashed to the ground. Loud-voiced
Science has told the people that characters acquired by individuals can under
no circumstances be transmitted to their descendants. It matters not if we live
good lives or bad, for, apart from direct disease or social degradation, we cannot
hand on to our descendants any vices, any talents or degeneracies that we may
have acquired during our lifetime.”

While we cannot quarrel with Wood Jones in his desire to find some
way to improve ethical standards, I think that we are justified in pointing
out how in this last quoted passage, he makes certain assumptions which
it would be extremely difficult to prove. As we know, the neo-Darwinian
theory is that evolutionary changes are due to adaptive selection of chance
variations due to mutations of genes ; and, further, Lamarckism, both
in its original form and in Wood Jones’ version of it, is dealing with somatic
characters. Now, even if we assume the validity of the Lamarckian
proposition, are we going to assume that because some modifications can
be transmitted, ethical modifications can be likewise transmitted ? It
appears to me that here Wood Jones is guilty of a deliberate confusion
of terms ; it does matter if we lead good lives or bad, not because we can
hand on our virtues and vices by heredity, but because man is a social
animal, and by living good lives we create an environment in which other
good lives can be lived. A sound ethical system is a social inheritance,
and not an individual somatic one.

That science may be related to ethics, and that the acceptance of the
orthodox biological theories is not incompatible with an attempt to devise
ethical systems, is shown in the address of a few months ago by Dr. Barnes,
Bishop of Birmingham (1950). Many of us who would agree with Dr.
Barnes’ ethical judgments would not agree with his theology. However,
that need not be a disturbing thought, for, as Leonard Huxley has written
(1920, p. 80) : —

“ It is alike interesting and satisfactory to reflect that practical morality
in civilized life is much the same for all earnest men, however they differ in
their theories as to the origin of moral ideas and the kind of motives and sanctions
to be insisted on for right action. It is true that the theologians and super-
naturalists have erected their scaffolding around the building of social and
human anatomy, vowing that it will not stand without. Yet it remains steady
when the scaffolding is warped by the winds of doctrine or uprooted by advancing
knowledge.”

FORM OR FUNCTION.

(b) In Physios.

The question of the relation of form and function is of interest and
importance in many fields other than the zoological — in botany, in geography,
in chemistry and physics. I have no doubt that all of you will have realised
this fully long ago. And now, having dealt at some length with some
more or less philosophical aspects of biology, I will beg your indulgence
for a few minutes longer, whilst I venture, in the words of Seneca, “ in
aliena castra transire, non tanquam transfuga sed tanquam explorator.”

With some trepidation, I propose to say a few words about the
philosophical implications of modern views of the form and function of
matter. The field of physics is particularly interesting, for here we find
that in the realm of micro-physics, i.e. intimate atomic “ structure,” the
physicist tells us that structure disappears in function, and mass has come
to be viewed as a form of energy ; and where previously we might have the
comfortable assurance that we could build a model of the atom, we now
find that we are not justified in such a procedure.

As you know, the Rutherford-Bohr atom succeeded the era of the
92 elements, each with its own sort of atom. In this new atom the picture
was simplified. Bohr’s theory of the structure of the atom was based
upon Rutherford’s conception of the atom as consisting of a small positive
charge of electricity (the nucleus) surrounded by a number of electrons.
Each atom could be regarded as a system composed of only two kinds of
constituents — electrons and protons. The electron has a negative charge
of electricity, the proton an equal positive charge ; the proton has about
1,850 times the mass of an electron. The nucleus is composed of a number
of protons and a smaller number of electrons. The nucleus is regarded
as taking the position of the sun, with electrons as planets revolving around
it, the number of planetary electrons being just sufficient to make the total
number of electrons (nuclear and non-nuclear) equal to the total number
of protons. Bohr assumed that the orbits were circular ; they were later
shown to be elliptical.

This atom had for a while a great success, as it explained many observed
facts almost perfectly. But after a reign of about twelve years it was
deposed in the mid-twenties in favour of the Heisenberg-Schrodinger
atom, which was a much more abstract and less imaginable affair. It is
still permissible to talk about electrons and protons just as we talk of sun-
rises and sunsets in spite of Copernicus ; but the ultimate truth is supposed
to be something quite different. New kinds of units — neutrons, positrons,
mesons and so on have been added, with such complexity of function that
it appears to be impossible to describe a model even as an extreme con-
cession to our desire for an imaginative picture of physical processes.

Now, Bohr’s atom was soon seen to present difficulties to classical
physics, according to which the energy of the atom ought gradually to
decrease, with the electron consequently assuming an orbit of ever decreasing
radius ; but such was not the case, for it seemed that there were only
certain admissible orbits and that while an electron, when it emits or absorbs
energy, can be said to “ jump ” from one orbit or energy level to another,
there can be no “ jump ” from an intermediate position. These admissible
orbits are those which possess angular momentum an integral multiple
h

of ~z~, h being Planck’s constant. “ Jumps ” may be “ up ” or “ down,’’

2tt

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

but only from the shown levels. Susan Stebbing (1944, p. 133) illustrates
the point by means of a diagram like this : —

Limiting energy level

The radical departure from classical physics can be illustrated from
radioactivity. Statistically we know in radioactivity just how many atoms
will decay, but we cannot give anything more than the probability for any
individual atom. The initial state does not “ determine ” the final state
in an individual case ; hence the final state is unpredictable. It is unfor-
tunate that this unpredictability has often been expressed by saying that
the electron is “ free to choose ” where it will jump. Such language is
wholly inappropriate and has led to much confusion in discussions concerning
the bearings of recent developments in physics upon the problem of free
will.

Put briefly, the problem is this — since every physical entity must
be measurable, how' is the position or the velocity of an electron to be
measured ? However, the means of measurement in this case affect the
phenomenon being investigated, so that in the case of an individual electron
it is not possible to ascertain with complete accuracy both the momentum
and the position of an electron : the more accurately the one is ascer-

tained, the more inaccurately is the other ascertained. The “ more ” is
formulated in Heisenberg’s “ Principle of Uncertainty,” which states that
the product of the uncertakriy of position and the uncertainty of momentum
is never less than Planck’s constant h, i.e. there is a theoretical limit to the
accuracy with which the state of a material system can be investigated.

Consequently it is not possible, even in principle, to know the initial
conditions in the case of quantum phenomena. The classical deterministic
system was based upon the assumption that the initial conditions can be
precisely known — which is true of macroscopic phenomena ; but no com-
pletely accurate predictions, except statistical, are theoretically possible
in the case of micro-physical systems.

Form or function.

The effects of this new concept on popular philosophy have been well
summarised by Crammer (ed.) (1950) : —

“ When in 1926 Heisenberg contributed his principle of Indeterminacy
to the development of the theory of quantum physics, a new era opened in
popular philosophy. What was intended to be a specialised theoretical con-
tribution to a restricted field of physics was quickly twisted by certain
popularisers of science and by delighted clergymen into meaning the general
overthrow of the laws of causality. The nineteenth century view of the Universe
as a great machine was suddenly seen to be hazy in fine detail ; there was still
room for human free will, for miracles and the intervention of the hand of God ;
for once, modern science had been compelled to grant living space to religion —
thus ran the new popular philosophy. It was a philosophy based on a mis-
understanding at its scientific root. Heisenberg had been concerned to point
out that we cannot by the nature of our methods measure simultaneously both
the velocity and the position of an atomic particle ; thereby stimulating closer
scientific inquiry of methodology in physics. But he said nothing about the
behaviour of particles being causeless ; this was added by the popularisers.”

However, to many it seemed that there was now a way of escape
from the nineteenth century nightmare of rigid determinism of all activity,
including mental activity and ethical judgment; i.e., the way was open
for some freedom of the will ; “ mind ” was restored to supremacy, material
objects could be regarded as “ abstraction,” and “ materialism ” was
supplanted by “ idealism ” ; 44 mind ” could be regarded as the creator.

What is the real situation ? Do the uncertain relations show that
there is anything indeterminate in Nature, or that science has now had to
become inaccurate ? Bertrand Russell (1946, p. 15) puts it simply : —

“ There are rules governing the changes that atoms undergo from one
energy-level to another, but these rules are not sufficient to determine which of
several possible things an individual atom will do. They do, however, suffice
to determine the average behaviour of a large number of atoms. The case is

analagous to throwing dice given a very great many atoms, all capable

of a certain definite set of transitions, we can tell, almost exactly, what pro-
portion will “ choose ” each possibility, though we cannot tell which will be
chosen by any particular atom. Consequently the behaviour of matter in bulk
is statistically deterministic, although each separate atom may make any one
of a certain definite set of transitions.”

Given that an experiment is repeated a great many times, then it is
possible to predict both the frequency of occurrence of the expected effects
and the deviation from these effects. There is nothing lawless in quantum
phenomena.

And on the question of whether physics is still deterministic, Russell
writes (p. 16) : —

“ The failure of determinism, where atomic occurrences are concerned,
has much less importance than is sometimes attributed to it. Except in a well-
equipped physical laboratory nothing can be discovered about the behaviour
of an individual atom or electron ; all the occurrences of which we are aware
in ordinary life involve many millions of atoms and are therefore just as
predictable as they used to be. It is true that the prediction is now only probable,
but the probability is so near to certainty that the element of doubt due to this
cause is very much less than that which will always be present owing to other
causes. . . For practical purposes, therefore, physics is still deterministic,
the only change is that the deterministic laws are all statistical.”

A further comment may be made. Although it is true that if the
rigid determinism of classical physics applied to human actions, these
would be completely unfree, still, as Susan Stebbing points out (1944,
p. 161), “it cannot be maintained that all that is required for human
freedom is some amount of uncertainty in the domain of microphysics.”
Can we really say that — “ physical events are not pre-determined, so that
it follows that human actions are not predetermined.” If we can say

20 PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

this, we are confronted by <e the difficulty of the ridiculously small amount
of indeterminacy that the measure of the uncertainty relations involves.
There is an appreciable degree of indeterminacj^ only in the case of electrons
and atoms ; human beings are polyatomic ; human actions take place in
the world of macroscopic bodies.” (p. 152).

Further, there is an immense cleavage between the behaviour of
inorganic bodies and the behaviour of a human being — for the latter is
able to tamper with the odds on atomic behaviour. This cleavage may
not do violence to physics, but it seems to leave the problem of free will just
where it was before physicists became indeterminists.

To my mind there is an insuperable difficulty to the transference
of quantum indeterminacy to human freedom of the will.

If there is a primary casualness in microphysics, when we transfer
this to the field of human action, surely we can only expect casual “ ethical ”
judgments — complete unpredictable caprice — which has nothing to do
with the character of the individual, that is, the very reverse of ethics.

We have already seen how Wood Jones has allowed his attitude to
genetics to be distorted by his views on the causation of the breakdown
in ethical standards ; and Susan Stebbing (1944, p. 166) has remarked
on how various thinkers have reacted to the contemporary situation in
physics each in conformity with his own philosophy of life : —

“ Planck is anxious to refute indeterminacy in physics in order to save
the dignity of man. Eddington is anxious to increase the amount of indeter-
minacy, recently introduced into physics, in order to safeguard our feeling of
responsibility. Sir Herbert Samuel is afraid lest the denial of determinism
should make man the sport of chance and lead to irresponsibility in action
and increase of unreason in politics and life.”

As I see it, the problem of k< the freedom of the will ” is, sociologically
at any rate, a false problem. The assumption of the reign of law is that
the law can be known, that the individual, unless disabled through infancy
or senility, disease or toxic substances, etc., is able to make a choice between
actions which we label “ good ” and “ bad.” If we do not accept this
assumption, we must agree that responsibility is an illusion ; if we do
accept it, we have two alternatives — either we must hold that physics
can be reconciled with the fact of responsibility and show how it can be
reconciled ; or we must agree that no reconciliation is required because
there is no conflict — this last is the practical solution so far accepted by
human society.

The false philosophical and ethical deductions from modern physics
are only part of a general picture of the “ retreat from reason ” of which
we see so many evidences to-day — man is being asked to save himself by
the acceptance of unreason, So well has the situation been expressed by
Susan Stebbing that I will close my address by quoting from the last paragraph
of her “ Philosophy and the Physicists ” (1944, p. 212) : —

“Is it not odd that men should come to this pass — that they look for
hope in physics and welcome, as some do, any indication of unreason in the
world ? Perhaps it seems less odd when we reflect upon the history of mankind,
the hopeless mess that we have made of human lives. Our greed, our stupidity
and lack of imagination, our apathy, these are the factors upon which the present
sorry state of the world is largely consequent. It is enough to fill us with despair.
Yet, despair need not be the last word. It lies within our power, if we so desire,
to make the familiar world we inhabit more worthy of habitation by beings
who aspire to be rational and are capable of love. Our limitation is due not to
ignorance, not to the ‘ blind forces of Nature,’ not to the astronomical
insignificance of our planet, but to the feebleness of our desires for good. This
limitation is not to be removed by the advance of physical knowledge, nor
should our hopes be placed in the researches of the physicist.”

FORM OR FUNCTION

REFERENCES.

Barnes, E. W. (Rt. Rev.) (1950). Science, Religion and Moral Judgements. Nature,
166, 455-7.

Bernal, J. D. (1939). The Social Function of Science. (Routledge, London).

Burnet, F. M. (1948). The Relation between Teaching and Research in the
Universities. Aust. J. Sci., 11, 5-7.

Crammer, J. L. (Ed.) (1950), Editorial, Science News No. 17. (Penguin Books).

Darlington, C. D. (1947). The Retreat from Science in Soviet Russia. The Nineteenth
Century and After, 1947.

Huxley, J. (1942). Evolution : The Modern Synthesis. (Allen & Unwin Ltd.,
London).

Huxley, L. (1920). Thomas Henry Huxley : A character sketch. (Watts & Co.,

London).

Huxley, T. H. (1895). Evolution and Ethics, and other Essays. (MacMillan & Co.,
London).

Kropotkin, P. (1902, 1919). Mutual Aid ; A Factor of Evolution. (Heinemann,
London).

McMurray, T. P. (1949). The Teaching of Anatomy to the Medical Student. Brit.
Med. J., 2, 510-514.

Medley, J. (1950). Farewell to Academe (The Sir Richard Stawell Oration). Med. J.
Aust., 2, pp. 949-954.

Romer, A. S. (1949). The Vertebrate Body. (W. B. Saunders Co., Philadelphia &
London).

Russell, B. (1946). Mind and Matter in Modem Science. The Rationalist Annual.
(C. A. Watts & Co., London).

Steering, L. Susan (1944). Philosophy and the Physicists. (Pelican Books).

Straus, W. L. (1950). Review in Quart. Rev. Bio1., 25, 95-6.

Wheeler, W. M. (1923). The Dry-Rot of our Academic Biology. Science, 57, 61-71.
Wigglesworth, V. B. (1934). Insect Physiology. (Methuen & Co., London).

Wood Jones, F. (1942). Design and Purpose. (Kegan Paul, Trench, Trubner & Co.,
London).

(1943). Habit and Heritage. (Kegan Paul, Trench, Trubner & Co.,

London).

Young, J. Z. (1950). The Life of Vertebrates. (Oxford University Press, London).

Zuckerman, S. (1947). Revision of the Anatomical Curriculum at Birmingham
University. Lancet, 2, 395-397.

SOME QUEENSLAND BLATTIDAE (ORTHOPTERA).

Vol. LXIII. No. 2.

STUDIES OF THE LIFE HISTORIES OF SOME
QUEENSLAND BLATTIDAE (ORTHOPTERA).

Part 1 . The Domestic Species.

By Pauline Pope, Queensland Institute of Medical Research.
(With 9 Text-figures.)

(Received 2nd April, 1951; issued separately, 6th July, 1953.)

INTRODUCTION.

Interest in Blattidae was aroused by finding two species naturally
infected with Salmonella during an epidemic of gastro- enteritis in Brisbane
in 1947 (Mackerras and Mackerras, 1948), and subsequently demonstrating
that artificially infected cockroaches might remain carriers for six weeks
(Mackerras and Pope, 1948). It was then realised that, although many
studies of domestic blattids have been made in other parts of the world,
we could find no comprehensive account of any species in Australia.
Answers to the questions “ How long do egg-capsules take to hatch ? ”,
and “ How long may adults live ? ” were necessary in planning a campaign
of control.

Descriptions of the domestic species are scattered through the literature
and are not always readily obtained. Brief systematic descriptions,
together with figures, are therefore mcluded to enable those interested to
identify the various species at any stage. An account of the life history
of each species is given. While many records of the life histories of
Periplaneta americana, Blattella germanica and Supella supellectilium have
been published, references to P. australasiae and Nauphoeta cinerea are
relatively scanty. None was found concerning P. ignota.

METHODS.

Most of the domestic cockroaches were easy to breed in laboratory
colonies. Some colonies were kept at room temperature and others in a
heated cupboard maintaining a temperature ranging from 73° to 84°F.
(22-8°-28'9°C). The graphs (text-fig. 1) show the monthly means of the
percentage of humidity and the temperatures in the cupboard and the
room.

Egg-capsules were isolated as soon as they were found and all nymphs
hatching from one egg-capsule were set up as a colony. Notes were made
on the incubation period, the duration of nymphal period, the appearance
of the first adult of each sex, the first egg-capsule produced and the total
length of life. In addition single females were isolated with one or more
males to obtain information about their egg-laying capacity.

The cockroach colonies were usually set up in dressing jars, measuring
6" x 6", and several layers of muslin were used as covers for the tops. The
bottom of the jar was covered with sterile sand ; pieces of bark and filter
paper gave extra surface area. Water was supplied in small dressing

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

1948 1949 1950

J JASON DJ FMAMJ JASOND JFMAMJ

Y FU.-HMgl

U EW

Text-fig. 1. Above. Four graphs showing the monthly means of the percentage
of humidity (at 9 a.m.) and the temperature of the heated cupboard (broken line),
the room (full line). Below. A diagram of the egg-to-egg cycles of domestic species
of Blattidae.

Stippled area

Egg incubation period.

Black area

Hatching to first female adult in a
(Appearance of first male marked by an

colony.

arrow).

Blank area

Preoviposition period.

Marks colonies kept in the heated cupboard.

I.-II. P. australasiae,
supellectilium.

III.-VI. B. germanica, VII. -VIII. N. cinerea,

IX.-X.

SOME QUEENSLAND BLATTIDAE (ORTHOPTERA) .

jars, 1" x 2", filled with wet cotton wool. Food consisted of a mixture of
dried milk, dried yeast, and cracked corn or wheat. Cake, “ Farex,”
bran and fresh apple were also given from time to time.

At first ether was used as an anaesthetic when it was necessary to sex
adults, or to transfer a colony to a clean jar. Later carbon dioxide was
used and found more satisfactory.

It was very difficult to maintain the right humidity for egg- capsules
during incubation. When the humidity was low, they tended to dry out
and when it was high, they became mouldy. The best results were
obtained by isolating them in sterile test tubes tightly stoppered with
cotton wool.

Descriptions have been taken from five insects, since pinned specimens
frequently become greasy and the distinctive patterns lost. In all species
variations in colour intensity are commonly seen.

A. DOMESTIC SPECIES OF PERIPLANETA.

The genus Periplaneta Burmeister (subfamily Blattinae) is represented
in Brisbane by three domestic species, australasiae , americana and ignota.
As far as I have been able to observe australasiae is more usual in dwellings
and americana in sewers and manholes. The most uncommon is ignota.

Their fully developed wings enable them to fly easily into houses
on summer nights and they can run very fast. They are particularly
averse to daylight, and in three years observation not once were any
Periplanetas seen mating in laboratory colonies. These species always
ate their own exuviae and dead, and very often their own egg capsules.

Adults of these domestic species are large, brown cockroaches, usually
about one to one and a half inches long and having the following characters
in common. Light brown head with darker vertex, white ocelliform spots,
interocular width less than inter antennal, long dark brown antennae,
yellowish- brown legs with brown spines, posterior metatarsus longer than
the other tarsal segments combined, small pulvilli, arolia present, anterior
part of wing yellowish-brown, anal area colourless with brown veins,
brown cerci considerably exceeding the supra-anal lamina. The male
subgenital plate bears a pair of unsegmented styles, and that of the female
is modified to a bivalvular structure (Text-fig. 3j).

The larger nymphs resemble the adults in general conformation. Styles
are present in nymphs of both sexes until the final moult when the female
loses them.

1. PERIPLANETA AUSTRALASIAE (Fabricius, 1775).

(a) Description of Stages.

adult (Text-fig. 2 AI). Pale yellow pronotum with dark brown
margins (considerably wider posteriorly than anteriorly), and with two
very dark brown maculae (often fused). Brown tegmina with yellow
humeral streaks. Yellowish-brown abdominal tergites and sternites,

darkening considerably towards margins and also apex of abdomen ; 7th
tergite backwardly produced. Dark brown cerci. The genital plates are
shown in text-fig. 3.

Total length* : $ 26-35 mm., $ 30-35 mm.

Tegmina length : $ 23-28 mm., $ 22-25 mm.

Pronotum width : $ 8-9 mm., $ 9-11 mm.

* The total length was measured from the vertex to the tip of the supra-anal plate.

5 mm

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

Text-Fig. 2.

A. P. austral asiae ; B. P. americana ; C. P. ignota ; I. male adult ; II. large
nymph ; III. newly hatched nymph ; IV. egg capsule.

SOME QUEENSLAND BLATTIDAE (ORTHOPTERA) .

Text-Fig. 3.

Male supra-anal plate : a, australasiae ; b, americana ; c, ignota. Male subgenital
plate : d, australasiae ; e, americana ; f, ignota. Dorsal view of tip of abdomen of
female : g, australasiae ; h, americana ; i, ignota. Ventral view of tip of female abdomen :
j, typical Periplaneta. Posterior tarsi of newly hatched nymphs : k, australasiae ;
1, americana ; m, ignota.

Text-Fig. 4.

Newly hatched nymphs (mounted specimens), a, australasiae ; b, americana ;

c, ignota.

labge nymph (Text-fig. 2 All). May be distinguished by the
following characters : — yellow pronotum with dark margins, considerably
wider posteriorly than anteriorly and with two large, dark brown maculae
and a brown median smudge wlr'ch extends to the tip of the abdomen.
Mesonotum and metanotum yellowish, with dark brown posterior margins
and with translucent wing pads developing in the larger nymphs.
Abdominal tergites brown medially, with pale spots ringed with darker
brown laterally on second to fifth and on seventh segments. Sternites
yellowish-brown. Arolia present.

28 PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

newly hatched nymph (Text-figs. 2 AIII and 4a). Head dark,
basal segments of antennae pale, four or five apical segments white,
remainder black. Dorsal surface dark brown to black with a white band
on mesonotum and another on the second abdominal tergite. The latter
is interrupted by a median black mark. Legs dark brown, tarsi pale.
Arolia present (Text-fig. 3k). Total length, 4-4*5 mm.; antennae length,
4*5 mm.

egg-capsule (Text-fig. 2 AIV). Dark brown, containing 24-26 eggs
side by side in 2 parallel rows. Length, 10-11 mm.; depth, 5 mm.

(6) Life History.

(i.) incubation period. Egg-capsules deposited from October to
February (i.e. during summer) hatched in 7 to 8 weeks, those laid in March
in 11 to 13 weeks, in April (one record) 23 weeks, in August about 10 weeks
and in September 8 to 9 weeks. We have no records of egg-capsules laid
during May to July. Those incubated in the warmed cupboard hatched
on the average in 53 days.

(ii.) duration of nymphal development. There was very great
variation in the rate of nymphal development. Although exposed to the
same climatic conditions, and having abundant supply of food, a lag of from
one to nine months was observed between the appearance of the first and
last adult in nymphs derived from a single egg- capsule. At room
temperature in summer the shortest period was 156 days for a male and
170 days for a female. There was almost invariably a considerable loss
by death between hatching and reaching adulthood, but this could not
be calculated at all accurately as dead nymphs (and exuviae) were almost
always eaten. Table I. gives the information obtained from some colonies
of P. australasiae.

TABLE I.

Duration of Nymphal Development in P. australasiae.

Colony

Number.

Site.

Date of Hatching.

Period (in Days) from Hatching
to Appearance of Adult.

Male.

Female.

Final.

200

Room

16 Feb

316

338

31 Mar

310

330

596

21 Sept.

156

170

374

106

99 • •

25 Nov.

309

335

410

110

26 Nov.

308

334

381

1 Dec.

286

337

143

„ • •

15 Dec. . .

268

301

342

Cupboard

26 May

195

239

285

„

21 July

184

223

236

19 Oct.

140

167

172

99 • • • •

9 Nov.

134

192

272

151

29 Dec. . .

142

198

373

(iii.) egg-laying capacity. At room temperature females produced
their first egg-capsule 15 to 48 days after their final moult, adult males
being already present in the colony. The average period observed was
21 days. In the warmed cupboard the range was 11 to 24 days, average
16 days. The minimum interval between ovipositions was 2 days and the
maximum 40 days. The productivity of seme individual females is set
out in Table II.

SOME QUEENSLAND BLATTIDAE (ORTHOPTERA) .

TABLE II.

The Egg-laying Capacity op Some P. australasiae Females.

Colony Number.

Site.

Total Number
of Egg-
Capsules
Produced.

Reproductive

Period

(Days).

Average

Interval

between

Ovipositions

(Days).

260

Room

251

109

119

Average

160

68a

Cupboard

136

141

154

267

108

175

164

Average

99 • •

147

The reproductive period referred to in Table II. is the interval between
the deposition of the first and last egg capsule by a female. In the
laboratory colonies the last few egg- capsules produced by a female, before
she died, seldom hatched and usually were smaller than the earlier ones.

In the laboratory colonies, australasiae females always endeavoured
to conceal their egg- capsules with sand particles, or minute pieces of filter
paper securely glued in place by a secretion from the mouth. Three days
was the longest time a female was seen carrying an egg capsule. The usual
time taken to produce and deposit one was 24 hours. It is produced
and carried with the serrated ridge dorsally.

Roughly the egg-to-egg cycle of australasiae covers 350-400 days at
room temperature, and 250-300 days in the warmed cupboard. The cycle
is so long that it cannot avoid the influence of the cold winter snap, and
consequently at least one period is considerably lengthened.

(iv.) longevity. The longest periods observed were 937 days for a
male and 897 days (i.e. about two and a-half years) for a female, both were
kept at room temperature. The average life-span for both sexes is about
18 months. In the warmed cupboard, under conditions similar to
continuous summer, cockroaches naturally died sooner. Table III. gives
the information obtained from some colonies.

TABLE III.

Longevity of some P. australasiae Adults.

Hatching to Death (Days).

Colony

Number.

Site.

Males (28).

Females (16).

Max.

Min.

Mean.

Max.

Min.

Mean.

Room

937

392

735 (7)

897

660

748 (3)

„

751

427

590 (5)

643

381

502 (7)

17a

Cupboard. .

284

284 (1)

645

645 (1)

30a

99 • • • •

666

666 (1)

533

533 (1)

713

470

557 (4)

427

359

393 (2)

141

99 • • • •

687

303

455 (9)

524

524 (1)

653

653 (1)

476

476 (1)

Numbers in brackets indicate the number of individuals observed.

30 PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

2. PERIPLANETA AMERICANA (Linnaeus, 1758).

(a) Description of Stages.

adult (Text-fig. 2 BX). Pale yellow pronotum with brown margins,
wider posteriorly than anteriorly, and with two light brown maculae (often
fused). Concolorous brown tegmina. Brown abdominal tergites and
sternites, 7th tergite not backwardly produced. Light brown cerci. Small
arolia present. The genital plates are shown in text-fig. 3.

Total length : 33-45 mm., $ 31-42 mm.

Tegmina length : $ 25-33 mm., $ 23-29 mm.

Pronotum width : 10-11-5 mm., § 11-13 mm.

large nymph (Text-fig. 2 BII). Almost uniformly brown, without
the pale abdominal spots of the other two species. Pronotum with two
slightly darker brown fused maculae, and a darker brown posterior margin.
Abdominal tergites with darker posterior margins. Sternites brown,
small arolia present.

newly hatched nymph (Text-figs. 2 Bill and 4b). Greyish-brown
body darkening slightly towards apex of abdomen. Posterior margins of
tergites slightly darker in colour. Long almost translucent antennae.
Slightly darker cerci. Legs translucent, arolia absent (Text-fig. 31).
Body length, 4*5-5 mm.; antennae length, 7 mm.

egg-capsule (Text-fig. 2 BIV). Differs from that of australasiae in
its smaller size and consequently in its more rounded appearance. Contains
16-18 eggs. Length, 8-9 mm.; depth, 5 mm.

(b) Life History.

(i.) incubation period. Observations were made on about 70 egg-
capsules. Those laid from October to February (i.e. during summer)
hatched in about 7 weeks, those laid in March in 7 to 8 weeks, in April
10 to 11 weeks, in August in about 9 weeks, and in September in about
8 weeks. We have no records for eggs laid from May to July. The
maximum period observed was 99 days. In the warmed cupboard the
average incubation period was 50 days.

(ii.) duration of nymphal period. The rate of development varied
greatly, and the recorded lag between the first and last adult was even
greater than with P. australasiae , ranging from 3 to 11 months. In these
observations the shortest period for complete development was 320 days
for a male, and 287 days for a female at room temperature. The periods
observed in some colonies of americana are given in Table IV.

TABLE IV.

Duration of Nymphal Development in P. americana.

Duration (in Days) from Hatching

Colony

Number.

Site.

to Appearance of Adult.

Date of Hatching.

Male.

Female.

Final.

Room

2 Aug.

477

505

813

26 Oct

365

359

—

29 Oct

320

287

—

9 Nov.

451

379

714

134

„ • •

19 Nov

335

337

452

Cupboard

20 July

325

274

395

340

24 Oct

257

186

—

25 Oct

192

134

331

5 Nov. . .

160

172

300

SOME QUEENSLAND BLATTIDAE (ORTHOPTERA) .

(ill.) egg-laying capacity. The preoviposition period varied a good
deal with the season. In summer at room temperature and in the warmed
cupboard egg-laying usually began about 2 weeks after the female’s final
moult if males were already present in the colony. The minimum period
observed was 13 days.

In some colonies females appeared considerably sooner than males.
If left thus unmated, they occasionally laid a few egg-capsules, which
usually looked very abnormal and invariably failed to hatch.

Takahashi (1924) reported that americana can produce partheno-
genetically a few, usually about three, egg capsules during life ; the young
from these did not normally hatch, and those that did, failed to mature,
However, Griffiths and Tauber (1942a) reported virgin females reproducing
parthenogenetically. This phenomenon occurred in 9 capsules laid by 3
of a group of 25 unmafed females. As many as 13 nymphs emerged from
one capsule.

In an attempt to confirm these findings, 6 virgin females were kept
in solitary confinement for periods ranging from 2 to 7 months. Of these,
four laid no eggs, one laid one egg-capsule and another laid 2 egg-capsules.
These were all soft, pale, obviously abnormal capsules, which failed to
hatch.

Females usually carried their eggs for 1 or 2 days. Sometimes they
glued them to the glass jar, but quite frequently they deposited them
loosely in the sand or food, in contrast to P. australasiae, which almost
always went to considerable trouble to fasten their eggs securely and to
conceal them with debris.

In the warmed cupboard their reproductive capacity was at a maximum
2 to 3 months after reaching the adult stage. Females then produced as
many as 7 capsules per month. Thereafter, the number gradually
decreased, until, towards the end of life, they laid only 1 or 2 per month.
At room temperature much the same thing occurred, but an early winter
diapause was followed by renewed activity later in the year. For example
a pair was mated in December, egg laying began in January, and the following
monthly totals were recorded : — January (5), February (5), March (6),
April (2), May (0), June (0), July (3), August (3), September (4), October (5),
November (4), December (6), January (5), February (4), March (4), April (1).

It is interesting to note that in this, and in two other colonies, egg-
laying was resumed in July — actually the coldest month. The egg-laying
capacity of some individuals is set out in Table V.

TABLE V.

The Egg-laying Capacity of some P. americana Females.

Colony Number.

Site.

Total Egg-
Capsules.

Reproductive

Period

(Days).

Average

Interval

between

Ovipositions

(Days).

Room

448

437

232

134

”

699

Average

454

8*7

Cupboard

268

”

246

Average

257

7-5

32 PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

(iv.) longevity. The longest period recorded was 1,502 days for
a male, and 1217 days for a female kept at room temperature. The
corresponding figures for the warmed cupboard were 1233 and 754 days
respectively. Table VI. gives the periods observed in some americana
colonies.

TABLE VI.

Longevity of some P. americana Adults.

Colony

Number.

Site.

Hatching to Death (Days).

Males (22).

Females (15).

Max.

Min.

Mean.

Max.

Min.

Mean.

Room

1,449

1,023

1,251 (3)

984

984 (1)

1,429

1,403

1,416 (2)

839

839 (1)

99 • •

1,194

942

1,068 (2)

693

693 (1)

941

885

910 (3)

—

134

1,502

1,329

1,415 (2)

1,217

1,217 (1)

Cupboard

1,233

841

1,004 (4)

645

533

607 (4)

99 • •

794

497

655 (4)

754

528

629 (5)

1,033

1,033 (1)

469

469 (1)

99 • •

998

998 (1)

546

546 (1)

3. PERIPLANETA IGNOTA Shaw, 1925.

(a) Description of Stages.

adult (Text-fig. 2 Cl). Light brown pronotum with pale yellow
anchor-shaped marking, and darker lateral and posterior margins.
Concolorous brown tegmina. Light brown abdominal tergites and sternites,
darkening laterally and towards apex of abdomen ; 7th tergite backwardly
produced ; dark brown cerci. The genital plates are shown in text-fig. 3.

Total length : (J 30-33 mm., $ 30-34 mm.

Tegmina length : $ 23-25 mm., $ 20-22 mm.

Pronotum width : 9-10 mm., $ 10-11 mm.

large nymph (Text-fig. 2 CII). Brown pronotum with fight median
marking. Light brown mesonotum and metanotum with dark posterior
margins. Dark brown abdominal tergites with pale lateral spots on 2nd
and 6th segments and a pair of very small pale spots on 3rd segment.
Light posterior margin on 7th tergite. Brown abdominal sternites
darkening laterally.

newly hatched nymph. (Text-figs. 2 CIII and 4c). Dark brown
head. Dark antennae with basal segments pale and five apical ones
white. Thoracic tergites brown, white transverse strip on mesonotum.
Yellow abdominal tergites and sternites, darkening considerably laterally.
Dark cerci. Pale legs with darker edges, arolia present (text-fig. 3m).
Length of body, 4*5-5 mm.; length of antennae, 5 mm.

egg-capsule (Text-fig. 2 CIV). Very similar to that of australasiae,
but each compartment slightly larger. Contains 22-24 eggs. Length,
12-13 mm.; depth, 5 mm.

SOME QUEENSLAND BLATTIDAE ( ORTHOPTERA ) .

(b) Life History.

This species is not present in Brisbane in as large numbers as
P. australasiae and P. americana, but it is found occasionally in dwellings.
Experiments were begun with a single female, which laid only one egg-
capsule in captivity. Additional colonies were set up when the progeny
of this female matured and commenced egg-laying. All were kept at room
temperature.

(i.) incubation period. During summer this varied from 49 to 61
days. Eggs laid in March required 11 weeks to hatch. Those laid in
April 13 weeks.

(ii.) duration of nymphal period. All colonies developed during
the summer months, the minimum periods observed were 1 10 days for a
male and 126 days for a female. The lag between the appearance of the
first and last adults varied from about 6 weeks to 7 months. Table VII.
gives the information obtained from 4 colonies of P. ignota.

TABLE VII.

Duration of Nymphal Development in P. ignota.

Colony

Number.

Date Hatched.

Duration (in Days) from Hatching to Adult.

Male.

Female.

Final.

31 Dec.

110

248

327

368

28 Nov

295

126

295

375

12 Dec

289

257

301

388

23 Dec.

270

158 1

311

(iii.) egg-laying capacity. From two colonies containing groups
of two and three females respectively, 59 and 89 egg- capsules were removed,
so on an average one female could produce nearly 30 egg capsules.
Unfortunately in each of these groups all the males died before the
reproductive period of the females had ended. Females were observed to
carry an egg- capsule for 1-2 days. They usually concealed their eggs in
the sand, glueing them firmly to the bottom of the jar. In this habit
and in the size of the egg-capsule they closely resembled P. australasiae.
Females usually deposited their first egg capsule about 12 days after
reaching adulthood. Thereafter they deposited them at intervals of about
a week.

(iv.) longevity. The periods observed in the original colony wTere
from 248 to 452 days, mean 390 days, for 7 males ; and 515 to 732 days,
mean 646 days, for 5 females.

B. OTHER DOMESTIC SPECIES.

Other domestic species of Blattidae occurring in Brisbane are
Blattella germanica (Linn.) and Supella supellectilium (Serv.), (subfamily
Pseudomopinae). Nauphoeta cinerea (Obv.) (subfamily Panchlorinae) has
been regarded as “ semi-domestic”, since it is found in outhouses and stores
rather than in dwellings.

Froggatt (1906) noted that B. germanica was first recorded in Australia
by Tepper (queried 1893, confirmed 1905).

B. germanica and S. supellectilium are small, light brown, very active
species about half an inch long. The former has two very distinctive,
longitudinal, black stripes on its pronotum ; the females carry their egg-
capsules attached to their bodies until the young are ready to hatch. The

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

latter has a plain brown pronotum with transparent edges ; the females
deposit their egg capsules soon after they are produced. N. cinerea is
a large, speckled, greyish -brown, sluggish species ; the females are
viviparous.

1. BLATTELLA GERMANICA (Linnaeus, 1767).

(a) Description of Stages.

Text-Fig. 5.

B. germanica. a, male adult ; b, female adult with wings clipped to show egg-capsule
c, large nymph ; d, and e, newly hatched nymph.

adult male (Text-fig. 5a). Light brown head with dark brown stripe
on anterior edge of vertex, inter ocular width slightly less than inter antennal,
white ocelliform spots and small brown median dot on frons, slightly
darker antennae. Light brown thoracic tergites with two longitudinal,
parallel, black ptripes. Light brown, coneolorcus tegmina ; pale, almost
colourless, wings with light brown costal and apical margins ; tegmina
and wings extending to tip of abdomen. First abdominal tergite rounded
and darker than rest, 2nd to 6th tergite yellowish, with dark brown
subapical markings laterally, and a pair of small transverse dark brown
medial basal markings, 7th and 8th tergites light brown with glandular
pockets,* 9th light brown with dark brown median marks ; 2nd to 6th

* Wille (1921) reported that copulation is preceded by a lengthy love-play in
which the female touches the dorsal glands of the male and then later licks them.

SOME QUEENSLAND BLATTIDAE ( ORTHOPTERA ) .

tergites with slight rounded backward projections, abdomen long and
slender in shape. Some dark thoracic sternal plates ; light brown abdominal
sternites. Light brown legs, coxae with slight black streaks, posterior
metatarsus longer than other tarsal joints together ; light brown cerci
considerably exceeding supra- anal lamina.

(b) female (Text-fig. 5b). Differs from the male in —

(i.) Genital plates (text-fig. 6). The emargination on the female
supra-anal plate varies in its size, and even may be completely
absent.

(ii.) General size. About the same length as the male, but abdomen
is much broader, reaching, and often extending beyond, the
tips of the tegmina (except when she is carrying an egg-capsule).

(iii.) Colour, (a) Generally much darker than the male ; ( b ) size

and intensity of dark marking on frons greater than in males.

Total length : £ 14 mm., $ 14 mm.

Tegmina length : £ 10 mm., $ 10*5 mm.

Pronotum width : £ 3-5 mm., $ 4 mm.

large nymph (Text-fig. 5c). Dark brown head, light brown vertex.

Yellow thoracic tergites with two wide longitudinal, dark brown stripes,
sometimes joining posteriorly on metanotum. Dark brown abdominal
tergites with pale dots on lateral margins becoming smaller towards 7th
tergite, and two median light brown dots on each tergite, most conspicuous
on 2nd, 3rd, 4th and 7th. Brown abdominal sternites darkening towards
apex and with pale lateral margins. Supra-anal plate mottled. Long
dark brown cerci always standing upright. Pale yellow legs, coxae darker
at base. Styles present in both sexes until the final nymphal moult, when
those of the female disappear and those of the males remain. (There is
great variation in the intensity of the light brown, median markings on
the abdominal tergites of large nymphs).

newly hatched nymph (Text-fig. 5d, e). Dark brown head and
body. Thoracic tergites with transparent edges, 2nd and 3rd with wide
yellow portion. Light brown, almost translucent legs with dark

streak on coxae ; arolia present. Dark brown antennae with pale basal
segments. Styles present in both sexes. Body length, 3 mm.; antennae
length, 3-5 mm.

egg-capsule (Text-fig. 5b). Ligho brown, slightly darker at outermost
end, and almost white at end attached to female’s body ; roughly
rectangular in shape ; usually contains 38-40 eggs. Length, 8 mm.;
depth, 3 mm.

(b) Life History.

The egg capsule of germcinica in its early period of formation was
noticed protruding from the abdomen of the female with the serrated
ridge in a dorsal position. When the capsule was almost fully extruded,
it rotated, and finally rested with the ridge lateral on the right side, when
looking at the female dorsally (See text-fig. 5b). It remained in this
position until the young were ready to hatch. Several instances showed
that the capsules would hatch after being removed from the female, only
if they were well matured, and not even slightly damaged when removed.
The abdomen of a female was always very contracted while she was
carrying a capsule.

36 PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

(i.) incubation period. At room temperature egg- capsules hatching
in midsummer were carried by a female for an average period of 24 days,
while in winter the time was 42 days. In the cupboard the average was
28 days.

(ii.) duration of nymphal development. The period of nymphal
development varied considerably according to the time of year when the
young were born. Those hatching in summer, or kept in the warmed
cupboard, developed very rapidly, reaching maturity in 7 to 10 weeks,
whereas those hatching in winter required over 4 months. Males usually
appeared first in a colony, but there was seldom a very great difference
between the sexes. The lag between the first and last adult, which was
such a marked feature in Periplaneta colonies seldom occurred, except in
colonies reaching maturity late in summer, when those nymphs which were
slowest in development were caught by the winter cold, and went on
developing slowly for several months after the first adults had appeared
(colony number 189). The results obtained in some colonies are set out
in Table VIII.

TABLE VIII.

Duration of Nymphal Development in B. germanica.

Colony

Number.

Site.

Date of Hatching.

Duration (in Days) from Hatching
to Appearance of Adult.

Male.

Female.

Final.

Room

3 Jan.

189

9 Feb

212

21 May

139

147

161

23 June

138

147

2 July

107

114

24 Sept.

126

26 Nov

132

6 Dec.

Cupboard

15 June

101

99 • • • •

30 July

121

30 Sept.

121

„ . .

29 Oct.

(iii.) number of moults. The determination of the number of moults
is difficult and tedious owing to the habit of most cockroaches eating their
exuviae. A special arrangement is necessary and it was attempted only
for B. germanica.

One or at most two newly-hatched nymphs were placed in a test tube,
fed on dried milk and yeast and watered by a 2 cc. ampoule full of water
plugged in the mouth of the test tube. They were kept in the warmed
cupboard. Daily examinations were made and the width of the head
capsule was measured at frequent intervals after anaesthetising the insect
with carbon dioxide. It was usually possible to predict when a nymph
was about to moult by the stretched appearance of the body.

Of a large number of nymphs set up in this way, 29 became adults,
8 females and 21 males. Eleven males became adult after the sixth moult,
but all the females and 10 of the males passed through seven moults before
reaching the adult stage.

(iv.) egg-laying capacity. Females usually produced 4 or 5 egg-
capsules.

SOME QUEENSLAND BLATTIDAE (ORTHOPTERA) .

(v.) longevity. B. germanica is a relatively short-lived cockroach,
living on an average about 9 or 10 months. Observations on 9 pairs which
were kept at room temperature, gave the following figures : — Males,
maximum 343, minimum 177, mean 260 days ; females, maximum 384,
minimum 202, mean 297 days.

The egg-to-egg cycle was taken from the day a female was carrying
a fully formed capsule until the day the first one of the progeny from that
egg case was carrying a capsule. In seven colonies kept at room
temperature the average duration of this cycle was 140 days. The
maximum was 206 days for one beginning in the early winter and the
minimum 88 in the early summer. In the controlled cupboard the cycle
varied from 109 to 162 days.

2. SUPELLA SUPELLECTILIUM (Serville. 1839).

Dr. Eland Shaw recorded the presence of this insect in Queensland
in 1924 when he wrote “ S. supellectilium (Serv.) occurs as a domestic
insect in company with the common Blattella germanica (Linn.) and seems
capable of even more rapid multiplication than that species. Its spread
over Australia generally is to be expected ” (Shaw, 1924). In 1925 he

wrote “ when Supella supellectilium (Serv.) invades places

already occupied by Blattella germanica (L.), it tends to oust the latter.”
(Shaw, 1925, p. 205). This species is widespread in Brisbane at the
present time, but usually it is found only in small numbers. We always
had very great difficulty in establishing laboratory colonies, and supellectilium
proved to be the most delicate of all the domestic species. It seems
doubtful if it could oust B. germanica in Queensland.

The male is generally a pale yellowish colour with fully developed
wings extending beyond the tip of its narrow, pale abdomen. The female
is generally darker and much broader than the male ; her shorter wings
do not reach the tip of the abdomen. The nymphs of this species can run
fast and jump well.

Text-Fig. 6.

Genital Plates. B. germanica. a, male supra-anal ; b, male subgenital ; c, female
supra-anal. S. supellectilium. d, male supra-anal ; e, male subgenital ; f, female
supra-anal ; g, female subgenital ; h, 7th tergite of male.

38 PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

(a) Description of Stages.

adult (a) male (Text-fig. 7a). Yellow to yellowish-brown head
(sometimes with rather orange vertex), white ocelliform spots, brown dot
below each antennal socket, interocnlar width less than interantennal,
dark brown antennae. Pronotum brown over body with transparent anterior
and lateral edges, very narrow, dark margin all round and narrow yellow
median stripe. Pale yellowish brown tegmina with transparent humeral
area, slight paleness in base of dividing vein with effect of a darker brown
band just below ; wings colourless with slight brown marking at apex ;
tegmina and wings both extending beyond tip of abdomen. First five
abdominal tergites colourless, remainder yellow, 7th with characteristic
brown impressed area (text-fig. 6h) ; postero-lateral angles of tergites
not backwardly produced. Supra-anal plate with slight emargination
(text-fig. 6d). Some dark thoracic sternites, abdominal sternites

Text-Fig. 7.

S. supellectilium. a, male adult ; b, female adult ; c, large nymph ; d, egg capsule.

darkening to orange at apex ; general shape of abdomen slender. Subgenital
plate narrow, roughly triangular with lobe-like styles (text-fig. 6a). Pale
yellowish legs darkening to orange tarsi, posterior metatarsus longer than
other tarsal joints together, pulvilli small and inconspicuous, arolia present,
tarsal claws symmetrical. Pale yellow cerci darkening towards tip,
extending considerably beyond supra-anal lamina.

(b) female (Text-fig. 7b). Short, brown tegmina with transparent
humeral margin with two incomplete transverse colourless bands near
base ; tegmina and wings covering about three-fifths of abdomen.
Abdominal tergites yellow with brown markings laterally on third to fifth
segments, sixth brown, remainder with centre brown and lateral portions
pale. Some dark thoracic sternal plates, abdominal sternites orange,
darkening towards apex of abdomen. Supra-anal plate with small emargin-
ation (text-fig. 6f). Subgenital plate orange, ample, rounded (text-fig. 6g).

Total length : and $ 12-13 mm.

Tegmina length : $ 11-12 mm., $ 8-9 mm.

Pronotum width : and $ 3 *5-4 *5 mm.

LARGE NYMPH (Text-fig. 7c). Dark brown head, usually lighter above
antennal sockets, antennae with yellowish- brown base, darkening to black
tips. Pronotum dark brown over body with transparent anterior and
lateral edges and some variable light marking in centre ; very pale yellow
mesonotum with H-shaped dark area ; pale yellow metanotum with dark
posterior margin. First abdominal tergite very dark brown, others pale

SOME QUEENSLAND BLATTlDAE ( ORTHOPTERA ) .

yellow (2nd to 5th with dark brown, lateral marks). Some dark thoracic
sternal plates ; yellowish abdominal sternites. Supra-anal plate with
small lateral dark marks, rounded triangular shape, no emargination. Very
pale yellow coxae and femora, more orange tibiae and tarsi. Cerci pale at
base, dark at apex.

newly hatched nymph. General colouration light greyish-brown.
Pronotum with pale lateral margins, mesanotum with central third white,
sides and posterior margins light brown, metanotum mainly white with
a light greyish- brown area on each side near anterior margin and along
posterior margin. Tergites light greyish- brown, ventral surface pale.
Legs very pale yellow almost translucent. Antennae similar in colour to
body except for the third segment which is paler. Styles present in
both sexes. Arolia present. Length, 2-6 mm.; antennae, 4 mm.

egg-capsule (Text-fig. 7d). Light brown, concolorous. As eggs
mature a definite medio-lateral green tinge develops. Usually containing
18 eggs, range 16-20. Length, 5 mm.; depth, 3 mm.

(b) Life History.

S. supellectilium appeared very fond of the gum on the backs of labels
and consequently some were included in its regular diet.

(i.) incubation period. The incubation period varied from 63 to
156 days. Eggs laid in July did not hatch until October; August eggs
hatched in November (about 90 days); those laid in November hatched
in 63 days. Eggs laid in March at the end of summer did not hatch until
July and August (maximum period observed 156 days). In the warmed
cupboard thej^ hatched in 7 weeks.

(ii.) duration of nymphal development. The period of nymphal
development varied considerably with the season ; however, even in
mid-summer its development was slower than that of B. germanica. The
minimum period observed was 90 days for a male and 98 days for a female,
developing at room temperature in mid-summer, the maximum period
observed from hatching to the appearance of the first adults was 223 days
in a colony hatching in late summer. The results obtained from some
colonies are set out in Table IX.

TABLE IX.

Duration op Nymphal Development op S. supellectilium.

Colony

Number.

Site.

Date of Hatching.

Duration (in Days) from Hatching
to Appearance of Adult.

Male.

Female.

Final.

163

Room

5 Jan.

161

236

355

239

11 Mar

223

241

5 Nov.

117

225

22 Nov

217

240

Cupboard

1 Sept.

114

138

14 Sept.

121

153

23 Sept

124

148

249

2 Oct

150

136

255

4 Nov. . .

117

111

182

40 PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

(iii.) egg-laying capacity. The preoviposition period varied from
8 to 9 days in the warmed cupboard to 63 days for a pair mated at the
beginning of winter at room temperature. Females usually carried their
egg-capsules for 1 or 2 days, but occasionally were observed to carry them
for longer periods up to 8 days. The capsules were produced and carried
with the ridge dorsal. Females laid from 6 to 25 capsules, the average
for 7 pairs being 14. Isolated females laid eggs at intervals ranging from
3 to 13 days, the most frequently observed interval being 7 days.

In several instances unmated supellectilium females produced either
apparently normal egg cases that failed to hatch, or a mass of whitish
eggs not covered at all by a capsule. The latter soon dried and shrivelled
on coming in contact with the air.

(iv.) longevity. S. supellectilium has a rather longer life-span
than B. germanica, the average period being a little greater than one year.

It must be remembered that it was the least adaptable of the domestic
species. Many nymphs died before reaching maturity and it is possible
that under more favourable conditions, it would live longer than in our
laboratory colonies. The longevity of some adults is set out in Table X.

TABLE X.

The Longevity op some S. supellectilium Adults.

Sex.

Place Reared.

Longevity (Days).

Max.

Min.

1 Mean.

Male

Room

667

259

490 (6)

Cupboard

397

173

| 272 (7)

Female

Room

538

154

! 385 (6)

Cupboard

240

207

| 228 (3)

Figures in brackets indicate number of individuals.

3. NABPHOETA CINEREA (Olivier. 1788).

This is a “ semi-domestic ” species. Although adults have been
found in dwellings, there have been no reports of its breeding there.
Usually this species is associated with grain stores and fowl-feeding pens.
No record of the life history of this species was found in Australian
literature. It was first recorded in Australia by Shaw (1918), having-
been taken at various localities between Brisbane and Cairns.

Both sexes have fully developed wings, which fall just short of the tip
of the abdomen. They have short, stout legs, and cannot run as fast as
the domestic species. They have a more scuttling movement. The
male subgenital plate bears a pair of unsegmented styles. The female’s
body is larger than that of the male.

(a) Description of Stages.

adult (Text-fig. 8). Fawn head with dark brown interocular band,
light interantennal band, interocular width less than interantennal, white
ocelliform spots, pale palps. Medium brown pronotum with fawn lateral
margins and symmetrical fawn picturing, dark brown, longitudinal stripe
on lateral edge of pronotal disc. Medium brown tegmina speckled with
white, translucent fawn costal area ; colourless wings with medium brown
veins ; tegmina and wings reaching at least to 7th tergite. Abdominal
tergites medium brown with white markings ; 1st to 7th not backwardly

SOME QUEENSLAND BLATTIDAE (ORTHOPTERA) .

Text-Fig. 8.

N. cinerea. a, male adult ; b, large nymph ; c, newly hatched nymph (mounted

specimen).

Text-Fig. 9.

N. cinerea. Genital plates, a, male supra-anal ; b, male subgenital ; c, female supra-

anal ; d, female subgenital.

produced, 8th only slightly so, abdominal sternites medium brown with
a few white markings, darkening towards apex. Pale legs, femora unarmed
beneath, short tibiae, large pulvilli, large arolia, tarsal claws symmetrical.
Supra-anal lamina rounded, one median notch on posterior margin (text-
fig. 9a male, and 9c female) ; pale cerci only slightly exceeding this lamina.
Male subgenital plate with a pair of unsegmented styles (text-fig. 9b) ;
female subgenital plate ample, rounded (text-fig. 9d).

Total length : 27-29 mm., $ 29-31 mm.

Tegmina length : $ 16-17 mm., $ 18-20 mm.

Pronotum width : 7-8*5 mm., $ 9-10 mm.

large nymph (Text-fig. 8b). Medium brown, shiny head and body.
White ocelliform spots, eyes slightly reduced, interocular width about equal
to interantennal, brown antennae, paler clypeus. Thoracic tergites with
very dark brown lateral margins ; abdominal tergites with dark lateral

42 Proceedings of the royal society of Queensland.

and posterior margins, generally darkening towards apex of abdomen ;
only 8th slightly backwardly produced. Some dark thoracic sternal
plates ; anterior abdominal sternites with median yellow tinge. Lighter
brown legs with dark spines on short tibiae ; large arolia. Medium brown
cerci not extending beyond supra- anal lamina.

newly hatched nymph (Text-fig. 8c). Head with greyish-brown
vertex, becoming paler towards the clypeus, palps translucent, eyes
reduced, interocular space greater than inter-antennal space, slightly
darker antennae. Greyish-brown body ; abdominal tergites with slightly
darker posterior margins ; ventral surface of abdomen grey, darkening
laterally and towards its apex. Supra-anal plate produced, one median
notch ; short cerci not projecting beyond this plate. Styles present in
both sexes. Light brown, almost translucent legs, tarsal claws symmetrical,
large arolia. Body length, 5 mm.; pronotum width, 2 mm.

(b) Life History.

In the laboratory this species was found to be particularly fond of
cracked maize, consequently this formed the main part of its diet,
supplemented by dried milk, dried yeast and fresh apple. It will eat its
own exuviae and dead (sometimes even before specimens are dead).

Mating was never observed in this species, although sometimes the
males would be seen running around the jars with their backs arched and
their wings standing upright. It is presumed that mating occurs at night.
The females are viviparous. The eggs are formed side by side in a double
row into an egg mass similar to that of other species. N. cinerea females
retain their egg mass, covered by a soft, transparent membrane within
their bodies until the young are ready to hatch. The plane of the egg
mass is parallel to that of the female’s body.

(i.) incubation period. As mating was not observed, and no egg'
capsule is produced, the length of the period of gestation was never observed
accurately. The period of gestation covers the interval from the pairing
of a male and a female to the production of young. Possible periods of
gestation were calculated from the appearance of a newly moulted adult
female in a colony, where one or more adult males were already present,
to the production of the first batch of young. These periods varied from
50 to 196 days, mean 105 days, at room temperature, and from 44 to 102
days, mean 64 days, in the warmed cupboard.

(ii.) egg-laying capacity. One female can produce as many as
4 “ egg masses ” each containing 30-40 eggs, usually at intervals of about
2 months. Often an old female produced a “premature ” mass of creamish
eggs, which did not hatch. On coming in contact with the air, it
immediately hardened and shrivelled.

(iii.) duration of nymphal development. The rate of nymphal
development was surprisingly rapid for such a large, sluggish insect. The
shortest period was 107 days in a colony set up in early summer. Males
usually appeared first. The results observed in some colonies are set out
in Table XI.

(iv.) longevity. N . cinerea is a long-lived species. The maximum

life-span recorded was 1,185 days for a male, and 1,026 days for a female.
Table XII gives the periods observed for some adults.

The “ egg-to-egg ” cycle, measured from the production of one batch
of young until the latter produced their first young, varied from 295 to
481 days, mean 372 days at room temperature (6 observations); and 182
to 246 days, mean 221 days in the warmed cupboard (5 observations).

SOME QUEENSLAND BLATTIDAE ( ORTHOPTERA ) .

TABLE XI.

Duration of Nymphal Development of N. cinerea.

Colony

Number.

Site.

Date of Birth.

Birth to Appearance of Adult (Days).

Male.

Female.

Final.

161

Room

10 Jan.

241

254

336

190

„

7 Feb

211

226

322

„

1 Mar.

279

322

332

„

30 July

215

285

109

„

24 Nov.

119

141

407

107

25 Nov

161

279

—

124

30 Nov.

107

156

274

149

99 * *

20 Dec.

213

338

470

Cupboard

21 Jan.

112*

209

2 Feb

187

191

342

6 Apr.

189

202

261

108

19 No v

136

146

—

136

99 • • • •

8 Dec. . .

139

— ■

243

* Sex not recorded.

TABLE XII.

The Longevity of some N. cinerea Adults.

Sex.

Place Reared.

Longevity (Days).

Max.

Min.

Mean.

Male

Room

1,185

324

550 (10)

Cupboard

646

383

473 (3)

Female

Room

1,026

381

803 (3)

Cupboard

619

296

398 (10)

DISCUSSION.

In the accessible literature are descriptions of the biology and life
histories of various cosmopolitan domestic species of Blattidae. Blatta
orientalis Linn., Periplaneta americana (Linn.), P. australasiae (Fabr.),
Blatella germanica (Linn.), and Supella supellectilium (Serv.). All these
species, with the exception of B. orientalis*, are common in Brisbane at
present.

It is interesting to notice that the species which have become
domestic pests in Queensland are all cosmopolitan and have been introduced.
The possible exception is Periplaneta australasiae (Fabr.). The origin of
this insect is not at all clear. Froggatt (1906) recorded that it was then
rare near Sydney, N.S.W., and regarded it as remarkable that a species
not common in its own country had become a serious domestic pest when
introduced into America. There seems no doubt that it was widespread
in America at the beginning of this century. Marlatt (1902) reported that
it was the most abundant and troublesome species in Florida. Shaw (1925)
considered that Fabricius used australasiae to mean “of Southern Asia,”
since the term Australasia as applied to Australia and New Zealand was
not in use when he described the insect in 1775. Probably it is also an

* The specimens of Blatta orientalis in the Eland Shaw collection in the Queensland
Museum were obtained at Kadina, S.A.

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

introduced pest. P. ignota Shaw was first described in 1925 from specimens
collected in Queensland, Shaw (1925). It is probably an endemic species,
but it is not abundant enough to be regarded as a serious pest.

P. americana was studied by Haber (1920) in U.S.A., Takahashi (1924)
in Formosa, Fischer (1928) in German}^ Nigam (1933) in India, Klein (1933)
in Palestine, and Gould and Deay (1938), Rau (1940) and Griffiths and Tauber
(1942) all in U.S.A. Studies of B. orientalis were carried out by Miall and
Denny (1885) in England, Rau (1924) in U.S.A. , Zabinski (1929) in Europe
and Qadri (1938) in England. The development of B. germanica was
observed by Wille (1921) in Germany, and Woodruff' (1938) in U.S.A.
S. supellectilium was studied by Cottam (1922) in Khartoum, and Back
(1937) in U.S.A.

The works of Marlatt (1915), Haber (1919), and Laing (1921) included
the above species except S. supellectilium , while the later reports of Gould
and Deay (1938 to 1940), Gould (1941) and Metcalf and Flint (1939) covered
them all, and included some others. Laing carried out his studies in
Britain, and the others in America.

When allowance is made for climatic differences, our findings agree
fairly well with those of the authors quoted. Gould and Deay (1938)
found the incubation period of P. americana to vary from 35 to 100 days, the
average for over 400 egg- capsules being 55 days. They found an average
preoviposition period of 13*4 days, and that one female could produce 59
eggs at an average interval of 5-9 days. The nymphal period varied from
285 to 616 days, average 409 days at a temperature range of 68° to 82° F.,
with relative humidity ranging from 27 to 61%. These authors noted that
the males of P. americana were usually longer in reaching the adult stage
than the females. This also occurred in our colonies. With P. ignota
(in the limited number of colonies observed) and with P. australasiae the
reverse was usually true. Gould and Deay (1938) record a maximum
life span for P. americana of 913 days, but Griffiths and Tauber (1942)
report that the life-span may exceed 1,200 days. A maximum of 1,502
days is recorded here.

B. germanica has probably been studied more intensively than any
other species. Gould (1941) records an egg-laying capacity of 5 capsules
per female, a figure which agrees well with my results. Laing (1921),
recorded a maximum of 7 egg-capsules per female. However, it seems
clear that this species lays relatively few egg- capsules ; its abundance
is due to its rapid development, to the large number of nymphs (38-40)
produced from each capsule, and to the greater protection from both enemies
and desiccation afforded to the eggs by being carried by the female during
incubation.

8. supellectilium produces a small egg-capsule usually containing
only 15-18 nymphs. The incubation period is a relatively long one and
nymphal development is slow. These factors probably contribute to its
scarcity in comparison with B. germanica. Gould and Deay (1940) gave
a good account of its life history in America. They found an incubation
period of 90 days at 73°F. and 49 days at 82°F. Nymphal develop-
ment required 161 days at room temperature, and 92 days at 84°F. Females
produced about 15 egg-capsules containing about 13 nymphs. The
minimum incubation period recorded by Cottam (1922) working in Khartoum
was 33 days.

SOME QUEENSLAND BLATTIDAE (ORTHOPTERA) . 45

Illingworth (1941) studied N. cinerea and recorded its viviparous
nature, finding 28-40 young produced at each birth. He also noted its
association with poultry food sheds in Honolulu.

SUMMARY.

The domestic species of Blattidae occurring in Brisbane are Periplaneta
australasiae (Fabr.), P. americana (Linn.), P. ignota Shaw, Blattella
germanica (Linn.), and Supella supellectilium (Serv.). Nauphoeta cinerea
(Oliv.) is a “ semi -domestic.” The Periplanetas deposit their capsules
within a day or two of formation. S. supellectilium carries them for 1 to 8
days, and B. germanica carries them until the eggs are ready to hatch.
N. cinerea is viviparous.

The maximum number of oviposit ions recorded was : — P. australasiae
(31), P. americana (68), P. ignota (30), B. germanica (5), 8 . supellectilium.
(25) and N. cinerea (4).

The usual numbers of eggs in a capsule were : — P. australasiae (26),
P. americana (16), P. ignota (24), B. germanica (40), 8. supellectilium (18)
and N. cinerea (40).

The rate of development of all stages was greatly influenced by
temperature, but there was also considerable variation in the rate "of nym-
phal development among siblings.

The incubation periods varied from 39 to 160 days for P. australasiae ;
39 to 99 days for P. americana ; 49 to 91 days for P. ignota ; 24 to 42
days for B. germanica ; 63 to 156 days for S. supellectilium.

The nymphal periods varied as follows : — P. australasiae 134-596
days, P. americana 134-813 days, P. ignota 110-327 days, B. germanica
49-212 days, S. supellectilium 90-355 days, N. cinerea 107-470 days.

The maximum life-spans recorded were : — P. australasiae 937 days,
P. americana 1,502 days, P. ignota 732 days, B. germanica 384 days,
S. supellectilium 667 days, N. cinerea 1,185 days.

The egg-to-egg cycles of the Periplanetas and N. cinerea cover about
a whole year. Two to three generations of B. germanica could be bred
each year and about two of S. supellectilium.

ACKNOWLEDGEMENTS.

I wish to thank Dr. M. J. Mackerras and Mr. R. Domrow of this
Institute for their great help with this work and in preparing the paper
for publication, and Mr. G. Mack of the Queensland Museum for permission
to study the Eland Shaw Collection.

REFERENCES.

Back, E. A. (1937). Proc. ent. Soc. Washington, 39, 207-213.

Cottam, R. (1922). Ent. mo. Mag., London, 58, 156-158.

Fischer, O. (1928). Mitt. Naturf. Ges. Bern., 1927 : V.-VII.

Froggatt, W. W. (1906). Agric. Gaz. N.S.W., 17, 440-447.

Gould, G. E. and Deay, H. E. (1938). Ann. ent. Soc. Amer., 31, 489-498.

(1938). Proc. Indiana Acad. Sci., 47, 281-284.

(1940). Bull. Indiana Agric. Exp. Sta., No. 451.

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

Gould, G. E. (1941). Proc. Indiana Acad. Sci., 50, 242-248.

Griffiths, J. T., Junr. and Tauber, O. E. (1942 a). Physiol. Zool., 15, 196-209.

(1942 b.) J. New York ent. Soc.,

50, 263-272.

Haber, V. E. (1919). Minnesota Agric. Exp. Sta. Bull., 186.

(1920). Ent. News, 31, 190-193.

Illingworth, J. E. (1942). Proc. Hawaiian ent. Soc., 11, 169-170.

Klein, H. Z. (1933). Z. Wiss Zool., 144, 102-122. Extract in Rev. Appl. Ent. (B),
21, 251. 1934.

Laing, F. (1921). Brit. Mus. Nat. Hist., Econ. Series No. 12.

Mackerras, M. J. and Mackerras, I. M. (1948). Aust. J. Sci., 10, 115.
Mackerras, I. M. and Pope, P. (1948). Aust. J. exp. Biol. med. Sci., 26, 465-470.
Marlatt, C. L. (1902). U.S. Dept. Agric., Divn. Ent., Circular 51 (2nd Series).
(1915). U.S. Dept. Agric. Farmers’ Bull., 658.

Metcalf, C. L. and Flint, W. (1939). Destructive and Useful Insects, McGraw
Hill, New York (2nd Edition).

Miall, L. C. and Denny, A. Y. (1886). The structure and life-history of the
cockroach ( Periplaneta orientalis). London, Lovell Reeve & Co.

Nigam, L. N. (1933). Ind. J. agric. Sci., 3, 530.

Qadri, M. A. H. (1938). Bull. Ent. Res., 28, 263-276.

Rau, P. (1924). Trans. Acad. Sci. St. Louis , 25, 57-79.

Rau, P. (1940). Ent. News. 51, 121-124, 151-155, 186-188, 223-227, 273 -278.
Shaw, E. (1918). Mem. Q’land Mus., 6, 151-167.

. (1924). Qld. Nat., 4, 115.

(1925). Proc. Linn. Soc. N.S.W., 50, 171-213.

Takahashi, R. (1924). L. Dobutsugaku Zasshi (Zool. Mag.) Tokyo, 36, 215-230.

Extract in Rev. Appl. Ent. (B), 12, 155-,1924.

Wille, J. (1921). Monographien zur angewandten Entomologie, No. 5, Berlin, 140

pp.

Woodruff, L. C. (1938). J. exp. Zool., 79, 145-167.

Zabinski, J. (1929). J. exp. Biol., 6, 360-386.

Vol. LXIII. No. 3.

STUDIES OF THE LIFE HISTORIES OF SOME
QUEENSLAND BLATTIDAE (ORTHOPTERA) .
Part 2. Some Native Species.

By Pauline Pope, Queensland Institute of Medical Research.

(With 3 Text-figures and Plate I.)

{Received 2nd April, 1951 ; issued separately §th July , 1953.)

INTRODUCTION.

While engaged on a study of the domestic species of cockroaches
common in Brisbane, interest was aroused in the numerous native species,
none of which appear to have been studied previously. Some of these bred
well in captivity, others proved less adaptable. Among the former were
species of the genus Methana (subfamily Blattinae) and some species of
Ellipsidiori , and Megamareta (belonging to the subfamily Ectobiinae).
Descriptions of the different stages and accounts of the life history of three
species of Methana , two of Ellipsidion and one each of Balta and Megamareta ;
are given. One new species of Methana is described.

A. THREE SPECIES OF METHANA STAL.

The three species of Methana studied were M. curvigera (Walk.), M.
marginalis (Sauss.) and M. caneae n. sp.

Most species of Methana have fully developed tegmina and wings
extending beyond the tip of the abdomen, but in several they are abbreviated.
This genus has been recorded from Australia, New Guinea and Borneo,
and is characterised by the supra-anal lamina in the male being quadrate,
margins not serrate, and in the female triangular, apex emarginate ;
pronotum anteriorly parabolic, posteriorly very obtusely angled ; posterior
metatarsus about equal in length to remaining joints, biseriately spined
beneath, its pulvillus apical ; remaining tarsal segments with large pulvilli,
not spined beneath ; tarsal claws asymmetrical. The females have the
typical blattine bivalvular type of subgenital plate and that of the male
bears a pair of unsegmented styles. The cerci are long and acuminate.

The favourite haunt of these species is under the loose bark of trees
or logs. Many specimens of curvigera were found in wattle trees. In the
strong sunlight they hid in curled up leaves. Their egg cases were found
attached to the underside of loose bark or leaves.

In laboratory colonies they were easily bred. Leaves and pieces of
bark were added to their rearing jars. They usually endeavoured to conceal
their egg-capsules with sand, food particles, or minute pieces of chewed
bark. Their regular diet was the same as that of the domestic species
(Pope, 1953). All the native species were reared at room temperature.

1. METHANA CURVIGERA (Walker, 1868).

Tepper (1893) cites Walker’s habitat of this species as Moreton Bay,
Queensland. Specimens have been collected at Fraser Island and at Mary-
borough on the adjacent mainland.

48 PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

This pale species has quite distinctive markings in black and reddish-
brown. The wide, pale or transparent margin around the whole insect is
a striking feature of both adults and nymphs. It adapted itself very well
to our laboratory conditions and was always very lively.

Text-fig. 1. A. M. curvigera (Walk.), B. M. marginalis (Sauss.). I, male adult;
II, large nymph ; III, newly hatched nymph (mounted specimen) ; IV, egg-capsule.
Adults, large nymphs and egg-capsules are drawn to same scale. Newly hatched
nymphs are greatly enlarged.

(a) Description of Stages.

adult (Text-fig. 1 AI). Head pale cream with dark, transverse
interocular band on vertex and pale inconspicuous ocelliform spots ; inter-
ocular width less than interantennal. Long, light brown antennae. Pronotum
pale, translucent, with dark brown f) -shaped band around lateral and
anterior portions of disc, the band being sometimes interrupted in the
mid line anteriorly. Left tegmen with wide colourless band along anterior
margin to apex, remainder reddish-brown. Right tegmen similar except
that the portion overlapped by the left tegmen is paler. Wings transparent
except for white markings in the radial areas and a brownish suffusion
basally over the branches of the median vein. Abdominal tergites pale,
developing medium brown tinges and dark brown lateral marks towards

SOME QUEENSLAND BLATTIDAE (oRTMOPTERA) .

Text-fig. 2. Genital plates of three species of Methana. Male supra-anal : a,
curvigera : b, marginalis : c , caneae. Male subgenital : d, curvigera : e, marginalis :
f, caneae. Female supra-anal : g, curvigera : h, marginalis : i, caneae.

7th, which is sharply backwardly produced; 8th to 10th pale, last with
small dark marks. Abdominal sternites reddish-brown darkening laterally,
but extreme lateral margins pale. Legs pale, dark spines, slight darkening
at tip of hind tibia, large arolia. Cerci pale yellowish -brown.

Total length : $ 22-23 mm., $ 20-24 mm.

Tegmina length : £ 17-18 mm., $ 16-18 mm.

Pronotum width : £ 7 *5-8-5 mm., $ 8-8*5 mm.

large nymph (Text-fig. 1 All). Thoracic tergites pale, almost trans-
parent laterally, outline of body marked by brownish-black line. Posterior
margins of pronotum and mesonotum dark brown, trace on metanotum.
Pronotum narrowly edged with brown. Dorsum of abdomen with broad
white lateral and posterior margins. Anterior tergites pale in centre,
becoming reddish-brown then black sub-laterally and posteriorly.
Abdominal sternites reddish-brown darkening laterally, but with extreme
lateral margins white.

newly hatched nymph (Text-fig. 1 AIIX). Yellow head, dark vertex,
black antennae. Yellow thoracic tergites, black line marking outline of
body, translucent edges. First abdominal tergite yellow, 2nd to 6th dark
laterally and posteriorly, the dark posterior band widening towards apex,
remainder yellow. Abdominal sternites brownish-yellow. Tarsi and tibiae
with slight apical darkening, arolia present. Body length : 3*5-4 mm.
Antennae length : 6 mm.

egg capsule (Text-fig. I AIV). Orange, shiny, mediolateral brown
dots (sometimes smudges) giving squared effect. Usually contains 12 eggs,
range 8-14. Length : 6-7 mm. Depth : 4 mm.

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

(b) Life History.

(i.) the incubation pertod varied from about 5 weeks in midsummer
to 8-9 weeks in midwinter.

(ii.) nymphal development. Males usually appeared first in a colony,
but sometimes males and females appeared simultaneously. The minimum
period observed was 176 days. The results obtained in 5 colonies are set
out in Table I.

TABLE I.

Duration of Nymphal Development of M. curvigera.

Colony

Number.

Date of Hatching.

Duration (in Days) from Hatching
to Appearance of Adult.

Male.

Female.

Final.

253

28 Mar.

246

259

266

297

27 May

181

228

317

21 June

176

203

310

27 June

193

190

—

316

7 July

180

235

(iii.) egg-laying. Preoviposition periods from 14 to 29 days were
recorded. One female kept with several males produced 16 egg- capsules
at intervals of about 8 days. The female produced the egg- capsule with
the serrated ridge dorsal, she usually carried it for one or two days and then
fastened it to bark or concealed it in the food.

(iv.) longevity. The total life span of males ranged from 404 to 505
days, mean 451 days (7 observations). For females the range was 264 to
515 days, mean 419 days (8 observations).

2. METHANA MARGIN ALIS (Saussure, 1884).

This large, brown species with fully developed tegmina and wings
in both sexes has a flavid margin on the anterior and lateral margins of
the pronotum, extending to the radial margin of the tegmina.

Tepper (1893) gives Walker’s habitat of this species as “Queensland,
West Australia.” It has been reported from North Queensland. Our
specimens were taken in South Queensland. It has sometimes been reported
as entering houses, but all our specimens were collected in the field.

(a) Description of Stages.

adult (Text-fig. 1 BI). Light coloured head, dark vertical marking
on irons not joining dark transverse bar on vertex, interocular width con-
siderably less than in ter antennal, white ocelliform spots, long brown
antennae. Rich shiny reddish- brown pronotum and tegmina, white band
around anterior and lateral edges of pronotum, white humeral streak on
tegmina extending beyond the level of the anal area. That portion of
right tegmen which is overlapped by the left is distinctly paler than the
remainder. Anterior part of wings light brown, posterior part transparent
with brown axillary veins. Abdominal tergites medium brown darkening
laterally and posteriorly and towards apex of abdomen, 2nd to 4th with
pale basal spots on the lateral margins, 3rd to 7th backwardly produced,
most conspicuously so in 6th and 7th. Abdominal sternites dark reddish-
brown, shiny, 1st to 3rd with pale lateral markings. Legs pale with brown

SOME QUEENSLAND BLATTIDAB (ORTHOPTERA) . 51

edges and spines. Hind tibiae brown. Cerci brown, considerably exceeding
supra-anal lamina. The genital plates of both sexes are shown in text-
fig. 2. Measurements of both sexes : —

Total length : 25-29 mm.

Tegmina length : 20-23 mm.

Pronotum width : 10-13 mm.

LARGE NYMPH (Text-fig. 1BII). Thoracic tergites reddish-brown
with white lateral margins joining anteriorly on pronotum, blackish wing
pads. Abdominal tergites reddish-brown with black posterior margins',
2nd to 7th backwardly produced, 2nd to 5th with light lateral edges ;
abdominal sternites light reddish-brown with pale markings laterally and
very narrow dark posterior margins.

newly hatched nymph (Text-fig. 1 Bill). Face dark brown, vertex
yellow, orange antennal sockets, black antennae with paler bases. Light
brown thoracic tergites with translucent edges and the dark outline of the
body visible. Slightly darker brown abdominal tergites with lateral
darkening, 6th to 7th with dark posterior margins. Abdominal sternites
light brown with lateral darkening. Dark brownish-black legs with orange
spines, large arolia. Cerci yellow at base, tip black. Total length : 4-5
mm. Antennae length: 7-5 mm.

egg-capsule (Text-fig. 1 BIV). Very large, dull orange brown colour
at sides ; dark brown, flat base ; serrated ridge. Usually contains 26 eggs,
range 24 to 30. Length: 11-13 mm. Depth: 4-5 mm.

(b) Life History.

(i.) the incubation period varied from about 5 weeks in midsummer
to 8 weeks in midwinter. The number of nymphs derived from one egg-
capsule varied from 12 to 26, average 18.

(ii.) nymphal development seemed less affected by temperature
than might be expected. Nymphs born in late summer and developing
during winter reached maturity in approximately the same time as early
summer nymphs. The results of observations on six colonies are set out
in Table II.

TABLE II.

Duration of Nymphal Development in M. marginalis.

Colony

Number.

Date of Hatching.

Duration (in Days) from Hatching to Adult.

Male.

Female.

Final.

237

8 Mar.

231

260

379

352

14 Nov.

256

239

274

358

1,7 Nov

250

243

258

361

21 Nov

183

246

294

372

12 Dec.

225

270

309

384

19 Dec.

221

241

—

(iii.) egg-laying. The minimum preoviposition period observed was

10 days, but in most colonies it was much longer, ranging from 3 to 6 weeks.
The female of a pair of adults, which were captured in the field, deposited

11 egg-capsules in 57 days, i.e. approximately one every 5 days. Observa-
tions on 2 isolated pairs were made. The females deposited 9 and 16 egg-
capsules respectively, at intervals ranging from 5 to 14 days. A female
usually carried her large egg- capsule for 1 to 2 days. It was usually fastened
to bark and covered lightly with debris.

52 PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

(iv.) longevity. Six males were observed to live from 339 to 473
days, mean 390 days ; five females lived from 275 to 589 days, mean 453
days. These figures refer to the total life-span, i.e. hatching until death.

3. METHANA CANEAE n. sp.

An undescribed species was found on Fraser Island, Queensland.
Specimens were obtained from under loose bark on a dead, upright tree.

It differs from marginalis (Sauss.) in (a) its smaller size, (b) the
abbreviated tegmina and wings, and (c) wider interocular space ; and
from parva Shaw in (a) the flavid humeral streak extending beyond the
level of the anal area, (b) the form of subgenital plate of the male, and (c)
vertical band on frons not joining that on vertex. This species has been
named in honour of Miss Helen Cane of the Division of Entomology,
Commonwealth Scientific and Industrial Research Organisation, Canberra,
who is at present working on a systematic review of Australian Blattidae
and to whom we are indebted for assistance in identifying native species.

(a) Description of Stages.

adult (Plate I, fig. 1. b, c). Pale head, dark vertical bar on frons
not joining dark transverse band on vertex, very inconspicuous ocelliform
spots ; interocular width only slightly less than interantennal width and
approximately twice ocular depth. Pronotum shiny, dark brown with a
wide white band around the anterior and lateral margins and a very narrow
dark line on extreme edge. Reddish-brown tegmina with pale humeral
streak fading away just before it reaches the apex ; portion of right tegmen
which fs overlapped by left is distinctly paler than the remainder ; wings
with anterior portion brown and posterior portion colourless with brown
axillary veins ; tegmina and wings extending to 6th abdominal tergite.
Abdominal tergites dark brown, 2nd to 5th with pale spots laterally, 2nd
to 7th backwardly produced. Abdominal sternites reddish-brown darkening
laterally and posteriorly. Dark cerci exceeding supra-anal lamina. Pale
coxae with dark stripe, pale femora, brown tibiae, dark tarsi ; tarsal claws
very asymmetrical. The genital plates of both sexes are shown in text-
fig. 2.

Total length : 19-22 min., $ 20-24 mm.

Tegmina length : ^ 11*5-13 mm., $ 12-5-13 mm.

Pronotum width : 8-9 mm., $ 8-5-9 mm.

large nymph (Plate I, fig. la). Light brown head, brown band on
front joining that on vertex, light brown antennae. Light, reddish-brown
thoracic tergites with wide, translucent margins, which are narrowly edged
with brown. First abdominal tergite reddish-brown, 2nd to 5th reddish-
brown in centre, dark brown laterally, margins of 2nd to 5th pale, remaining
segments dark brown. Abdominal sternites reddish-brown’ small dark
lateral dots. Pale coxae with dark streaks, tibiae light brown, darkening
towards tarsi. Reddish-brown cerci.

newly hatched nymph (Plate I, fig. le). Yellowish head, dark
brown antennae. Yellow thoracic tergites, 2nd to 3rd with pale orange
posterior margins. Abdominal tergites yellow with orange posterior
margins, 6th with brown lateral dots. Orange abdominal sternites darkening
very slightly laterally. Yellow coxae, legs darkening slightly from femora
to tibiae, arolia present. Yellow cerci with dark tips. Total length : 3
mm. Antennae : 5 mm.

SOME QUEENSLAND BLATTIDAE (ORTHOPTERA) . 5$

egg-capsule (Plate I, fig. Id). Orange-yellow with irregular medio-
lateral brown markings, flat base giving squarish effect. Serrated ridge.
Usually contains 22 eggs. Length : 9-10 mm. Depth : 4 mm.

distribution. Fraser Island, Queensland, (Feb.), type locality.

taxonomic notes. Holotype male, allotype female, morphotype
nymphs and egg- capsule, bred in laboratory from adults collected at Fraser
Island, Q. ; in collection of the Division of Entomology, C.S.I.R.O., Canberra.

(b) Life History.

(i.) the incubation period varied from 4J-5 weeks in midsummer
to 7-8 weeks in winter. The number of nymphs hatching from one egg-
capsule varied from 12 to 22, average 16.

(ii.) nymph a l development. The most rapid development occurred
in a colony set up in early spring, adults of both sexes appearing by the
20th week. The results obtained from six colonies are set out in Table III.

TABLE III.

Duration of Nymph al Development in M. caneae.

Colony

Number.

Date of Hatching.

Duration (in Days) from Hatching to Adult.

Male.

Female.

Final.

252

28 Mar.

200

205

270

11 Apr.

203

213

227

278

19 Apr.

224

202

—

284

22 Apr.

195

—

290

29 Apr.

188

200

—

325

25 Aug.

139

132

— .

(iii.) egg-laying. The preoviposition period varied from 12 to 26
days. The female usually carried the egg- capsule for 1-2 days and fastened
it to bark. Two colonies containing 2 and 3 pairs were kept under observa-
tion. The former produced 69, the latter 79 egg-capsules. The period of
reproductive activity lasted about one year. Egg-laying continued
throughout the winter, though there was definite falling off in production
in April, May and June. In each of these colonies females survived con-
siderably longer than the males, and the last few egg-capsules produced
were infertile.

(iv.) longevity. The total life-span of 5 males ranged from 344 to
568 days (mean 472 days), that of 5 females ranged from 562 to 702 days
(mean 622 days).

B. SOME ECTOBIINE SPECIES.

The species belonging to the subfamily Ectobiinae which were studied
were Ellipsidion affine Hebard, E. australe Sauss., Balta scripta (Shelford)
and Megamareta verticalis Hebard. E. affine and E. australe were collected
in trees near Brisbane, B. scripta on Fraser Island and near Brisbane, and
M. verticalis at Gordonvale (North Queensland).

All these species have the following characters in common : — (i) female
subgenital plate not of the valvular (blattine) type; (ii) anterior femora
with antero- ventral margin lacking heavy spines before the row of piliform
spines and with one or two terminal spines; (iii) median and posterior
femora with ventral margins armed with spines; (iv) tarsal claws simple
and decidedly asymmetrical.

In Ellipsidion and Balta the tegminal discoidal sectors are oblique,
while in Megamareta they are longitudinal.

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

1. ELLIPSIDION AFFINE Hefeard, 1943.

E. affine is a small species which has a black abdomen, the lateral
and caudal margins of abdominal sternites being edged with white. These
are the markings characteristic of the genus. Both sexes have fully
developed tegmina and wings.

All the young hatching from one capsule were set up as one colony.
This proved rather awkward because in a colony there often appeared a
very great interval between the first moults to adult and the last.

Text-fig. 3. Genital plates. Ellipsidion affine : a, male supra-anal ; b, male
subgenital ; c, female, supra-anal ; d, female subgenital. Balta scripta : e, male supra-
anal; /, male subgenital; g, female supra-anal. Megamareta verticalis: h, male supra-
anal ; i, male subgenital ; j, female supra-anal.

(a) Description of Stages.

adult (Plate I, fig. 2 b). Black face, orange vertex, small pale orange
ocelliform spots, black antennae with wider and very hairy basal half and
orange distal half ; interocular width greater than interantennal. Orange

SOME QUEENSLAND BLATTIDAE (ORTHOPTERA) .

pronotum with translucent margins, darker over body, sometimes with
faint dark marks on pronotal disk. Orange tegmina with black bases
and slightly black apices, exposed part with checkered effect ; orange
wings with wide, black border extending from apical to anal area. Some
black thoracic sternal plates ; abdominal sternites black, 3rd to 6th with
definite white lateral and posterior margins, faintly marked on 2nd, 7th
white laterally, 8th and 9th wholly black. Orange cerci with black bases,
considerably exceeding supra-anal lamina. Black coxae with definite
white stripe on posterior margin, orange legs, apical halves of femora black,
tarsi dark (except for slight orange tinge on posterior metatarsus). The
genital plates of both sexes are shown in text -fig. 3. Measurements for
both sexes are :

Total length : 11-13*5 mm .

Tegmina length : 9*5-11*5 mm.

Pronotum width : 4*5-5 mm.

large nymph (Plate I, fig. 2 a). Black pronotal disk bordered laterally
with orange, white posterior margin, transparent lateral margin. Mesonotum
and metanotum black with orange translucent lateral margins and white
posterior margins. Abdominal tergites 1st and 2nd rounded, 3rd to 7th
with postero-lateral angles backwardly produced ; first five tergites with
white dots on posterior margin and a transverse white line just anterior to
the row of dots, 6th and 7th with posterior margin white and two median
white dots ; 8th and 9th black with white posterior margin ; 10th wholly
black. Some dark thoracic sternal plates ; abdominal sternites black,
1st five with white lateral and posterior margins, 6th with white lateral
margins. Black coxae with white stripe, orange trochanter, black femora
with orange tinges, tibiae orange in centre, remainder black, dark tarsi,
large arolia.

newly hatched nymph. Dark brown head, translucent antennae
darker at tips ; inter ocular width greater than inter antennal. Dark brown
pronotum ; golden brown mesonotum and metanotum ; all thoracic tergites
with translucent margins. Dark brown abdominal tergites. Medium
brown abdominal sternites. Golden-brown cerci. Dark brown legs with
transparent tarsi. Styles present in both sexes. Body length, 1*5-2 mm.;
antennae length, 1*5 mm.; pronotum width, 1 mm.

egg capsule (Plate I, fig. 2c). Cream with brown medio-lateral
dots, sometimes slightly darker below these ; dark brown base. Contains
30-32 eggs. Length, 6 mm.; depth, 2*5 mm.

(b) Life History.

(i.) incubation period. In the summer months the egg incubation
period was usually 25 days while in the winter it was twice as long.

(ii.) period of nymphal development. As our series began in the
middle of the summer season, the nymphs which did not reach maturity
quickly had a very long nymphal period extended by the winter snap. The
minimum period from hatching to the appearance of adults was 59 days,
the majority, however required 10 weeks to complete their development.
Nymphs hatched in February did not become adult until August, average
period 175 days. In most colonies there was a considerable lag between
the appearance of the first and last adults, for three colonies for which
records were kept, it varied from 5 to 35 weeks.

56 PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

(iii.) copulation. Apparently daylight does not disturb this species
very much, because the females could be seen depositing egg-cases, and
pairs could be seen preparing to copulate and actually copulating. On
several occasions the whole process was observed. The pair obviously
appeared interested in each other, and rushed backwards and forwards
frantically waving their antennae. The male walked around with his wings
upright and his body arched so that the tip dragged over whatever he crawled
on. At the same time the cerci pointed inwards and downwards together in
the same plane as the abdomen. The male stood right in front of the female
with the tip of his abdomen under her head. Then he poked his body
under hers, and, as the female crawled on to his back, she appeared to wipe
his tergites with her palps. When the tip of the male’s abdomen reached
that of the female he grasped her genitalia. Almost as soon as they con-
tacted they swung round end-to-end in a flashing movement and then
copulation took place. If the male failed to grasp the female, the pair
would break away and start again. From the literature it appears that
copulation in some cockroaches is a very speedy process. However, in
the laboratory we have observed pairs of E. affine in copulo for periods as
long as an hour. The female dragged the male in whatever direction she
chose while the pair was in copulation.

This species copulated very frequently. A short observation on one
pair is quite indicative.

Date.

Action of Pair.

6 XII. 48

Pair copulating

13 XII. 48

Female depositing

16 XII. 48

Pair copulating

21 XII. 48

Female depositing

22 XII. 48

Pair copulating

23 XII. 48

Female depositing

27 XII. 48

Female depositing

1 I. 49

Pair copulating

3 I. 49

Female depositing

6 I. 49

Female depositing

Five days after a pair matured they copulated, and 9 days after this
the female was carrying an egg case. Actually in our colonies the pre-
oviposition period varied from 14 to 30 days. The egg- capsule was produced
with the serrated ridge dorsal, it was usually only carried for a day and
then fastened on to a stem or the underside of a leaf.

(iv.) egg-laying capacity. The total egg-laying capacity was not
determined, but one female laid 8 egg- capsules in 6 weeks, the average
interval being 6 days, another laid 5 in 44 days. The egg-to-egg cycle is
measured from the deposition of one capsule through until the nymphs
hatching from it mature and deposit their first capsule. In the summer
season it was as short as 113 days, while cycles in the winter season were
extended to well over 200 days. It is possible for two generations of this
species to be bred each year.

2. ELLIPSIDION AUSTRALE (Saussure, 1864).

This species has the characteristic markings of the genus i.e. black
sternites edged with white. It is larger than E. affine, being about 19 mm.
long and 6 mm. wide (at level of pronotum). The general colouration is a
darker shade of orange and the black markings on the tegmina and wings
are more pronounced. The pronotum is black with a yellow margin. It
is an arboreal species like E. affine ; and the nymphal stages of these two
species are very similar. (Plate I, fig. 3 a-c).

SOME QUEENSLAND BLATTIDAE (ORTHOPTERA ) .

Life History.

The incubation period varied from 3-4 weeks in midsummer to 6-7
weeks in midwinter. The number of eggs per egg-capsule was usually 32.

The nymphal period for nymphs hatching in early summer was about
18 weeks, those hatching in later summer required 32 weeks.

The preoviposition period was about 3 weeks. Females produced
the egg- capsule with the serrated ridge dorsal and fastened it to bark or
leaves, but did not attempt to conceal it. One female adult, collected
in the field, laid 8 egg-capsules in 5 weeks, the average interval being 5
days.

Some insects were observed to live from 312 to 441 days, i.e. from
hatching to death.

3. BALTA SCRIPTA (Shelford, 1911).

Baita scripta is a small, greyish-brown speckled cockroach. It was a
very difficult species to handle in the laboratory. The adults were extremely
lively, and the newly hatched nymphs very small and practically colourless.
The small, light brown egg-cases were usually well concealed by the female
either in the food, or sand in the bottom of the jar, and they were always
extremely difficult to find.

(a) Description of Stages.

adult (Plate I, fig. 4 b). Light brown head; dark transverse inter-
ocular band with cream transverse band immediately below it, and another
dark transverse band immediately below this again ; interocular width
less than interantennal ; light brown antennae darkening towards tip.
Pronotum light brown with transparent lateral margins and a symmetric
design in dark brown lines and dots on the disk. Tegmina and wings fufiy
developed in both sexes. Tegmina with checkered effect, transparent edge
along lateral margin (when folded in repose). Wings with distal end of
costal veins thickened. Abdomen light brown. Light brown cerci pro-
jecting considerably beyond supra-anal lamina. Pale yellowish -brown
legs. The male subgenital plate is very asymmetrical (Text-fig. 3 f), that
of the female simple, ample and slightly emarginate on the free margin.
The supra-anal plates are shown in text-fig. 3 e, g.

Total length : ^10*5 min., $ 9*5 mm.

Tegmina length : 9 mm., 5 8 mm.

Pronotum width : $ 3*3 mm., $ 3*3 mm.

large nymph (Plate I, fig. 4 a). Pale head ; dark transverse inter-
ocular band with pale transverse band below it and another dark transverse
band below this, and then an interantennal transverse row of dots. Pro-
notum widest posteriorly, translucent lateral margins, pale pronotal disk
with symmetric design in dark brown lines and dots. Thoracic and
abdominal tergites pale, symmetrically marked with dark brown lines
and dots. Abdominal sternites pale with dark brown markings. Cerci
pale with dark brown marks, considerably exceeding supra-anal lamina.
Pale legs, dark stripe at base of coxae, slight darkening at base of spines.

newly hatched larva. Pale cream transparent head, body and
appendages ; large arolia. Total length : 1*3 mm. Pronotum width :
0*73 mm.

58 PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

EGG capsule (Plate I, fig. 4 c). Light brown, concolorous ; length of
serrated ridge greater than basal length. Usually contains about 16 eggs.
Length : 3 mm. Depth : 2 mm.

(b) Life History.

The incubation period was 28-30 days for eggs laid in January and
February. The nymphs energing from a capsule in July (that is, about
midwinter) matured 142-148 days later, while those emerging in February
matured in 230 days. The longevity of two males was 219 and 231, while
the intervals between their final moult and their death were 71 and 83
days. Another male lived 146 days as an adult. Two females which produced
7 and 8 egg capsules lived for 68 and 181 days respectively as adults. The
longevity of the first female was 216 days from hatching.

4. MEGAMARETA VERTICALS Hebard. 1943.

This is a relatively large, broad species (in comparison with other
Ectobiinae), uniformly pale yellow in colour. It adapted itself quite easily
to our laboratory conditions, and was probably the liveliest and most
prolific of the ectobiine species we bred.

(a) Description of Stages.

adult (Plate I, fig. 5b). Pale yellow head and body. Interocular
width less than interantennal ; white ocelliform spots ; pale antennae
darkening slightly towards tip ; darker brown vertical band on face, often
expanding near ocelliform spots to become almost T-shaped ; brown
interocular band. Pronotum with transparent lateral margins. Tegmina
with transparent humeral streak, wings with distal ends of costal veins
slightly thickened. Pale cerci projecting considerably beyond supra -anal
lamina. Legs uniformly pale yellow. The genital plates of both sexes
(except the female subgenital, which is simple and ample) are shown in
text-fig. 3 h-j. The measurements of both sexes are as follows : —

Total length : 13*5-15 mm .

Tegmina length : 12-13 mm.

Pronotum width : 5 mm.

large nymph (Plate I, fig. 5 a). Pale yellow head and body. Thoracic
tergites with broad transparent margins ; all with two small brown dots
posteriorly. Pale yellow abdomen ; 1st to 5th abdominal tergites with
row of small brown dots on posterior margin, 6th to 10th with small dark
markings on lateral margins. Pale cerci with slightly dark bases, con-
siderably exceeding supra-anal lamina. Pale, almost transparent legs.

newly hatched nymph. Very pale head and body with almost
transparent appendages. Very wide interocular space. Thoracic tergites
each with two dots posteriorly. Mottled cerci. Large arolia. Total length :
2 mm.

egg capsule (Plate I, fig. 5 c). Very dark brown, smooth, not shiny,
very narrow ; compartments do not show up distinctly ; wider at ridge
than base; usually contains 30-36 eggs. Length, 6-7 mm.; depth, 3 mm.

(b) Life History.

During midsummer the incubation period ranged from 34 to 38 days,
in early and late summer it covered 40 to 50 days and in midwinter 63 to
76 days.

Nymphs emerging in the early summer matured in 90 to 130 days.

SOME QUEENSLAND BLATTIDAE (OETHOPTEHA) .

The preoviposition period varied from 7 to 17 days. In a colony
containing four females kept under observation for 9 months over 70 egg-
capsules were produced. The female of the original pair was already an
adult of unknown age when our laboratory series began. She produced 10
egg-capsules in 84 days, at intervals ranging from 3 to 22 days. In the
gravid female the abdomen became distinctly green just before she produced
an egg-capsule.

The total life-span ranged from 257 to 408 days, mean 344 days, for
11 males ; and 265 to 424 days, mean 350 days, for 14 females.

Acknowledgments. — I wish to thank Miss H. Cane of the Division
of Entomology, C.S.I.R.O., Canberra, for identifying these native species ;
and Dr. M. J. Mackerras and Mr. R. Domrow of this Institute for assistance
in the work and in the preparation of the manuscript.

SUMMARY.

One new species, Methana caneae n. sp. is described together with its
life-history. Brief descriptions and life-histories are given of Methana
curvigera (Walk.), M. marginalis Stal, Ellipsidion affine Hebard, E. australe
(Sauss.), Balta scripta (Shelf ord) and Megamareta verticalis Hebard.

The periods observed for egg incubation, nymphal development and
maximum life span respectively are as follows : —

M . curvigera :
M. marginalis :
M . caneae :

E. affine :

E. australe :

B. scripta :

M. verticalis :

5 to 9 weeks ; 25 to 38 weeks ; 515 days.

5 to 8 weeks ; 26 to 44 weeks ; 589 days.

4J to 8 weeks ; 19 to 32 weeks ; 702 days.

3J to 7 weeks ; 8J to 45 weeks ; 315 days.

3 to 7 weeks ; 18 to 32 weeks ; 441 days.

4 weeks (summer) ; 20 to 33 weeks ; 230 days.

5 to 11 weeks ; 13 weeks (summer) ; 424 days.

REFERENCES.

Pope, P., (1953). Studies of the life histories of some Queensland Blattidae (Orthoptera),
Part 1. The domestic species. Proc. Roy. <Soc. Queensl., 63, 23-46.

Tepper, J. G. O., (1893). The Blattariae of Australia and Polynesia. Trans. R. Soc.
S. Aust., 17, 25-126.

EXPLANATION OF PLATE I.

Fig. 1. Methana caneae : a, large nymph; b, female adult; c, male adult;
d, egg-capsule; e, newly hatched nymph. Fig. 2. Ellipsidion affine. Fig. 3, E. australe.
Fig. 4. Balta scripta. Fig. 5. Megamareta verticalis : a, large nymph; b, adult; c,
egg-capsule. Figs. 1 a-d are to same scale ; scale beside 1 e is in mm. ; figs. 2-5 are
to the same scale.

5 mm

Pkoc. Roy. Soc. Q’land., Vol. LXIII., No. 3.

Plate I.

Vol. LXIII. No. 4.

PARASITES OF THE BANDICOOT, ISOODON

OBESULUS.

By I. M. Mackerras, M. J. Mackerras and D. F. San bars,* Queensland
Institute of Medical Research.

( Received 30 th July, 1951; issued separately 6th July, 1953.)

A considerable number of parasites from bandicoots in south-east
Queensland has been collected during the past two years. Several of the
records are new, and it has been thought worth while to bring them all
together in this preliminary list, even though some of the identifications
are incomplete.

We are indebted to Dr. E. H. Derrick of this Institute for referring
certain species to us.

PROTOZOA.

Trypanosoma sp. : Found in the blood. There are no previous records
in Australian marsupials, but a large trypanosome has been found
in the platypus in Tasmania.

Eaemogregarina ?peramelis Welsh and Dalyell : Described in 1910

from Perameles nasuta\ apparently this is the first record since
then. Our material does not agree completely with the original
description.

Theileria sp. : Species of Theileria are well known in Eutheria (T .

mutans was introduced into Australia in cattle) and have been
found in Monotremes as follows : — T. tachyglossi Priestley in
Tachyglossus aculeatus, and Theileria sp. in Ornithorhynchus
anatinus (Duncan, personal communication) ; they have not
previously been recorded from marsupials.

Klossiella sp. : Found in the kidney of bandicoots in Brisbane by

Derrick and Smith (personal communication).

Encephalitozoon sp. : This is an obscure genus which has not been

recorded previously from marsupials. An infection was found
first in mice inoculated from a bandicoot and later confirmed
by finding a single group of parasites in sections of the brain of
the bandicoot (E. H. Derrick, personal communication).

Sarcocystis sp. : Found in voluntary muscle of the body wall. This

genus was recorded by Bourne (1934) in Bettongia , but is not
otherwise known from marsupials.

PLATYHELMINTHES.

Trematoda :

Brachylaemus similis (S. J. Johnston), syn. Harmostomum simile.

Platynosomum Looss (new record). The recovery of Platynosomum
sp. appears to be the first record of a pancreatic fluke from an
indigenous Australian mammal.

* and Department of Social and Tropical Medicine, University of Queensland.

62 PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

Cestoda :

Hymenolepis peramelidarum Nybelin.

Linstowia echidnae (Thompson) (recorded by other workers but not
taken by us).

Linstowia semoni Zschokke (recorded by other workers but not taken
by us).

ACANTHOCEPHALA.

Gigantorhynchus semoni Linstow.

NEMATODA :

Capillariinae (new record). These very slender worms were found
embedded or half embedded in the oesophageal walls of the host.

Echinonema cincta (Linstow), syn. Hoplocephalus cinctus Linstow.
Filarinema peramelis T. H. Johnston and Mawson.

Subulura peramelis Baylis.

Trichuris peramelis Baylis.

Metastrongylidae. The first record is made of a lung worm
from an Australian marsupial.

Filarioidea. A new species of Dipetalonema was found in the sub-
cutaneous tissue. No microfilariae were found in the blood, but
on making sections of the skin they were found immediately
below the Malpighian layer. The finding of microfilariae in the
skin suggests relationship with Onchocerca , but the worm is
quite distinct morphologically. The life history is still unknown.
The intermediate host may be a biting insect, e.g. a mosquito
or sandfly, or it may be a mite.

ECTOPARASITES.

The ectoparasites of this bandicoot are quite numerous, the following
having been recorded either from Isoodon obesulus or I. macrourus. Dr.
E. H. Derrick and Mr. D. J. W. Smith included many of them in their
studies on Q fever, and published a list in the Annual Report of the Health
and Medical Services of the State of Queensland for 1937-38.

ACARINA.

Ticks. — Ixodes holocyclus Neumann ; I. tasmani Neumann ; /. fecialis
Warburton and Nuttall ; Haemaphysalis humerosa War burton
and Nuttall.

Mites. — Heterolaelaps antipodianum Hirst ; Mesolaelaps australiensis
Hirst ; M. anomalus Hirst ; Laelaps nuttalli Hirst ; Ascoschon-
gastia dasycerci (Hirst); A. cairnsensis (Womersley and Heaslip) ;
A. peramelis (Womersley); A. phascogale (Womersley and
Heaslip) ; Trombicula deliensis Walch ; T. minor Berlese ;
Guntherana bipygalis (Gunther) ; Bdellonyssus bursa (Berlese) ;
Listrophoridae, species not yet identified.

INSECTA.

Mallophaga. — Boopia sp.

Siphonaptera. — Acedestia chera Jordan ; Stephanocircus dasyuri Skuse ;
Pygiopsylla zethi Jordan and Rothschild ; P. congrua Jordan and
Rothchild ; P. hoplia Jordan and Rothchild ; Otenocephalides
canis (Curtis) ; C. fells (Bouche).

PARASITES OF THE BANDICOOT, ISOODON OBESULUS.

REFERENCES.

Bouene, G. (1934). — Sarcosporidia. Proc. R. Soc. W. Aust., 19, 1-8.

da Costa Lima, A. and Hathaway, C. R. (1946). — Pulgas. Monograf. Inst. Oswald.
Cruz, No. 4, 526 pp.

Johnston, T. H. (1916). — A census of the endoparasites recorded as occurring in
Queensland, arranged under their hosts. Proc. R. Soc. Q'land , 28, 31-79.

Peiestley, H. (1915). — Theileria tachyglossi (n. sp.) a blood parasite of Tachyglossus
aculeatua. Ann. troy. Med. Parasitol., 9, 233-238.

Welsh, D. A. and Dalyell, E. J. (1910). — Haemogregarina peramelia . a free
haemogregarine of an Australian bandicoot. J. Path. Bad., 14, 547-549.

Womeesley, H. (1937). — Studies in Australian Acarina Laelaptidae. Parasitol, 29,
530-538.

Womeesley, H. and Heaslip, W. G. (1943). — The Trombiculinae (Acarina) or itch-
mites of the Austro-Malayan and Oriental Regions. Trans. R. Soc. S. Aust.,
67, 68-142.

Young, M. R. (1939). — Helminth parasites of Australia. Imp. Bur. Agr. Parasitol,
(Helminthology). England, 145 pp.

Vol. LXIII. No, 5.

A STUDY OF DIPHYLLOBOTHRIiDAE
(CESTODA) FROM AUSTRALIAN HOSTS.

By Dorothea F. Sandars, Queensland Institute of Medical Research, and
Department of Social and Tropical Medicine, University of Queensland.

(With Plates II. and III.)

(. Received 2 6th November, 1951; issued separately 6th July, 1953.)

INTRODUCTION.

Following the recovery of numerous spargana from a local host, an
investigation was undertaken to identify the adult of these parasites and
determine whether laboratory animals would serve as “ reservoir ” hosts
for the spargana.

The author would like to express appreciation for the co-operation
given by Dr. M. J. Mackerras during this work ; to Mr. J. Thomson and
Dr. M. C. Bleakly who identified the copepods and frogs respectively,
and to Mr A. J. Bearup for many helpful suggestions and Mr. G. Thompson
who carried out some of the photographic work.

PREVIOUS RECORDS OF DIPHYLLOBOTHRIIDAE IN
AUSTRALIAN HOSTS.

The following records were found in a search of the a vailable literature ;
those marked with an asterisk were recorded by Young (1939).

adults : —

*Bothridium arcuatum Baird, 1865, in Python spilotes (N. S. Wales).
*Boihridium pythonis Blainville, 1824, in Python spilotes (N. S. Wa»les).

*Bothridium pythonis var. parva Johnston, 1913, in Varanus varius
(Queensl.).

Bothridium ornatum Maplestone and Southwell, 1923, in Python spilotes
var. variegatus (Queensl.).

*Bothriocephalus marginatus Krefft, 1871, in Macropus sp. (Queensl.).

*Diphyllobothrium latum (Linne, 1758) in Homo sapiens (Queensl.,
N. S. Wales, Viet., Tas.). (See Sandars, 1951).

DiphyUoboihrium latum (Linne, 1758) in Canis familiaris (N. S, Wales)
recorded by Gordon (1939). (See Sandars, 1951).

*Diphyllobothrium parvum (Stephens, 1908) in Homo sapiens (Tas.).

*Diphyllobothrium antarcticum (Bail'd, 1853) in “ Southern Seal ”
(Antarctic Ocean).

*Diphyllobothrium ardocephalinum Johnston, 1937, in Arctocephalus
forsteri. (S. Austr.).

*Diphyllobothrium decipiens (Diesing, 1850) in Dasyurus sp. (Austr.),
This record is probably incorrect. It appears as though it was
originally recorded from a domestic cat and not from the native
cat, Dasyurus sp. (See Cobbold 1879, p. 308).

66 PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

* Diphyllobothrium mansoni (Cobbold, 1882) in Felis domestica (Queensl.).

* Dibothriocephalus felis (Creplin, 1825) in Felis domestica (Queensl.,

N. S. Wales and Viet.).

Diphyllobothrium erinacei ( Rudolph i, 1819) in Vulpes vulpes (Viet.)
and in Canis familiaris (Viet.) recorded by Pullar (1946).

According to Neveu-Lemaire (1936), Diphyllobothrium decipiens ,
D. mansoni and Dibothriocephalus felis are probably synonyms of
Diphyllobothrium erinacei (Rudolphi, 1819).

Wardle, McLeod and Stewart (1947) claim that Diphyllobothrium
latum should be Dibothriocephalus latus Linnaeus 1758 ; Diphyllobothrium
arctocephalinum Johnston, 1937, should be Cordiocephalus arctocephalinus
(Johnston, 1937) ; and that decipiens, erinacei, felis and mansoni of
Diphyllobothrium should be included in the genus Spirometra (Mueller,
1937).

SPARGANA : —

Sparganum mansoni (Cobbold. 1883) three records in Homo sapiens*
(N. S. Wales).

Sparganum sp. in Homo sapiens * (N. S. Wales) ; Vulpes vulpes* (S.
Austr.) ; Dasyurus viverrinus* (N. S. Wales) ; Chlamydosaurus
kingii * (Queensl.) ; Dendrophis punctualatus * (Queensl.) ;

Demansia textilis * (Queensl.) ; Pseudechis australis* (Queensl.) ;
Pseudechis porphyriacus * (Queensl. and N. S. Wales) ; Python
spilotes* (N. S. Wales) ; Python spilotes var. variegatus* (N. S.
Wales and Queensl.) ; Varanus gouldii* (Queensl.) ; Varanus
varius* (Queensl.) ; Hyla aurea* (N. S. Wales and W. Austr.) ;
Hyla caerulea* (Queensl. and N. S. Wales) ; Thyrsites atun * (N.
S. Wales).

Bearup (1948) records the sparganum of Diphyllobothrium erinacei in
Acanthopis antarctica (Heathcote, near Sydney, N. S. Wales). He reared
the adults in experimental kittens and infected the following copepods
with procercoids : — Mesocy clops obsoletus (Koch), Cyclops australis (King)
and Leptocyclops sp., probably Leptocyclops agilis (Koch).

Pullar and McLennan (1949) record Sparganum sp. in the pig, Sus
scrofa (Viet.).

Recently spargana from “ wild ” domestic pigs from N. S. Wales have
been fed to both cats and dogs. Adult Diphyllobothrium erinacei were
recovered (Personal communication from Dr. H. McL. Gordon).

RECORDS OF NATURALLY INFECTED HOSTS.

Spargana have been recovered from the following hosts, which were
taken in the greater Brisbane area : —

(а) Natrix mairii Gray. Fresh- water Snake.

(б) Pseudechis porphyriacus Shaw. Red-bellied Black Snake.

In one specimen of Natrix mairii, spargana were found lying close to
the muscles on the dorsal side of the body cavity and smaller forms occurred
in the fat around the gut. Another specimen of Natrix mairii was very
heavily infected with spargana which occurred between the skin and body
wall, within the muscles of the body and throughout the body cavity ;
these spargana were most abundant in the middle third of the entire body
length (Plate II, figs. 1-3). From one Fresh-water Snake, over 300 spargana,
ranging in length from 5-213 mm., were collected.

A STUDY OF DIPHYLLOBOTHRIIDAE (CESTODA).

In Pseudechis porphyriacus, spargana with lengths between 5-40 mm*
were recovered from the peritoneum of the body cavity.

In Hyla caerulea, spargana were most commonly found between the
muscles of the inside of the thigh regions of either hind leg. They were
also recovered from between the muscles of the shoulder region.

EXPERIMENTAL.

(1) SPARGANA FROM NATRIX MAIRII.

On May 25th, 1950, spargana from the fresh- water snake (N atrix mairii)
were fed to two young cats, A and B ; each cat was given 6 spargana. One
sparganum from the same host was also fed to a. laboratory- bred white
rat. Twenty-three days later (June 17th), diphyllobothriid eggs, with
average measurements 57p, x 3 were recovered from the faeces of the
cats (Plate III, fig. 1). The faeces were washed thoroughly in tap water,
the eggs thus obtained being put into tap water in petri- dishes. One of
the developing larvae within an egg measured 32p, x 23/x, and each of the
six hooks present were of equal length, 1 1/x. On July 29th, some of the
eggs had hatched, and free swimming coracidia were observed. On the
same day, locally obtained copepods, Cyclops varicans Sars, were placed in
the petri-dish. On September 13th, the copepods were observed to be
infected with procercoid larval forms. (Plate III, fig. 2). The males, as is
usual, were observed to be infected, often heavily, with Diphyllobothrium
procercoids, while the females were not infected. Some infected copepods
were found dead on September 19th, and on examination they proved to
be very heavily parasitized with procercoids ; one examined contained
7 procercoids, 6 of them situated in the tail region.

On September 19th, two tadpoles of Hyla latopalmata (Gunther)
were introduced into a petri-dish of fresh water containing several
infected copepods. On November 11th the tadpoles were infected with
plerocercoids which were conspicuous, whitish structures lying just under
the skin of the host. They were found in various parts of the body and
tail on the dorsal and lateral surfaces of the host (Plate III, figs. 3, 4).
The growth of these tadpoles was obviously inhibited. In one, the dis-
tortion was very evident, especially in the region where the tail joins the
body (Plate III, fig. 4).

These spargana were fed, on November 27th, 1950, to various animals : —

(а) To a small frog, Hyla latopalmata (Gunther) bred in the laboratory
from a tadpole collected at Camp Mountain, near Brisbane. It was fed
one sparganum dissected from the infected tadpole. On May 7th, 1951, a
sparganum of increased size was recovered from between the muscles on
the inside of the thigh of a hind limb. The frog was killed by chloroform.
The sparganum had also been killed and had macerated very quickly.

(б) One sparganum was fed to a laboratory-bred mouse (Mus musculus
albus). On March 15th a sparganum of increased size was recovered from
between the muscles of the back, in the region behind the right fore-limb.
This was fed to an experimental cat. Soon afterwards this animal broke
its back and had to be destroyed. No tapeworm was recovered.

(i) One cat was fed 1 sparganum dissected from the tadpole. Sixteen

days later, on December 13th, a young specimen of Diphyllo-
bothrium erinacei was recovered.

(ii) One cat was fed a dead tadpole containing live spargana. On

December 24th, Diphyllobothrium eggs (average size 56/x x 33 ja)
were moderately abundant in the faeces of the cat.

68 PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

The white rat which was also fed a sparganum from Natrix mairii
on 25th May, 1950, was killed and examined on December 14th, and a
sparganum 120 mm. long was recovered from between the muscles of the
left thigh. This sparganum had increased considerably in length, having
been only 20-30 mm. long when taken from the fresh- water snake and fed to
the rat. It was then fed on December 14th to a cat, and on January 11th,
1951, Diphyllobothrium eggs (average size 63 /x x 30/x) were seen in the cat’s
faeces. On May 21st, one adult Diphyllobothrium , erinacei was recovered
from the intestine of the cat.

(2) SPARGANA FROM PSEUDECHIS PORPHYRIACUS.

Two host specimens were examined and in both, spargana were very
abundant. Spargana from one host were placed in a corked tube and kept
in a refrigerator until the following day. They were then removed and
shortly afterwards placed in 0*85% saline and were observed to be
alive and very active. Three of these spargana were fed to ar experimental
cat on October 15th, 1951 ; twelve days later (October 27th) the cat died
and no diphyllobothriids were recovered.

(3) SPARGANA FROM HYLA CAERULEA.

One of the most common frogs in the greater Brisbane area is Hyla
caerulea White, approximately one quarter of the population of which
appears to be infected with spargana. Two spargana recovered from
locally obtained Hyla caerulea were fed to an experimental cat on December
5th, 1950. A few diphyllobothriid eggs (average size 60 /x x 30/x) were
observed in the faeces on January 6th, 1951 ; on January 18th eggs, of
average size 62/x x 31/x, were abundant. The minimum time between the
feeding of spargana and the appearance of diphyllobothriid eggs in the
faeces of the host was 23 days.

Longevity of Diphyllobothrium erinacei.

Of the 2 cats, A and B, fed with spargana from Natrix mairii on May
25th, 1950 : —

(a) From cat A, 4 adult Diphyllobothrium erinacei were recovered from
the small intestine on November 3rd, 1950. They were all about 30 cm.
in length with a maximum width of 0-5 cm. The scolices were buried
fairly deeply in the intestinal mucosa.

(b) In the faeces of cat B, Diphyllobothrium eggs (average size, 62/x x
31/x) were still abundant on November 22nd, 1951. (In the same faeces
some eggs measured 52/x x 31/x). On November 28th, only a few Diphyllo-
bothrium eggs were present in the cat’s faeces and no eggs were recovered
on December 12th.

Variance in Ego Size of Diphyllobothrium erinacei.

Neveu-Lemaire (1936, p. 398) states that the size of the eggs of
Diphyllobothrium erinacei is very variable, measuring 52 /x to 76/x long, by
31/x to 44/x wide.

Eggs recovered from the several specimens of Diphyllobothrium erinacei
during the present work, varied considerably in size, the most usual egg
size being 62/x long, by 31/x wide.

DISCUSSION.

Bear up (1948) showed experimentally that spargana from Acanthopi
antarctica (Death Adder) were those of Diphyllobothrium erinacei. Since
spargana from Natrix mairii and Hyla caerulea have now also been shown

A STUDY OF DIPHYLLOBOTH RIID AE (CESTODA).

experimentally to be of Diphyllobothrium erinacei it seems that this tape-
worm is established, at least in eastern Australia, and that a wide range
of animals may serve as “ reservoir ” hosts for the sparganum stage in its
life history. According to Galliard and Ngu (1946) and Gan (1949) the
tadpole is an important host in which the plerocercoid stage develops
from the procercoid. Under natural conditions they must serve as effective
transmitters of spargana to other intermediate hosts, such as frogs, lizards,
snakes etc.

The first larval stage, the procercoid, was bred successfully by Bearup
(1948) in several species of fresh-water copepods from N. S. Wales, viz.
Mesocy clops obsoletus (Koch), Cyclops australis (King) and Leptocyclops
sp., probably Leptocyclops agilis (Koch). Galliard and Ngu (1946) note
that previous workers had infected successfully a number of different
species of Cyclops in other parts of the world ; they themselves infected
2 species, viz. Cyclops leuclcarti Claus syn. Mesocyclops obsoletus (Koch),
and another unidentified species. It has been shown experimentally that
Cyclops varicans Sars from Queensland also serves as a host for the procercoid
larval stage of Diphyllobothrium erinacei.

Bearup (1948) recorded the appearance of eggs of Diphyllobothrium
erinacei in the faeces of the kitten 45 days after the spargana had been
fed to it. In the present experiments eggs were recovered in abundance
as early as 23 days after spargana had been fed to a cat. This indicates
that the worms reach their adult stage in the final host in a period of just
over three weeks.

SUMMARY.

(i) A new host record is made for the sparganum of Diphyllobothrium
erinacei (Rudolphi), viz. Natrix mairii Gray (Brisbane, Queensl.).

(ii) Spargana from Natrix mairii and Hyla caerulea from the Brisbane
area were shown experimentally to be of Diphyllobothrium erinacei.

(iii) The copepod, Cyclops varicans Sars, was shown to act as a host
for the procercoid stage in the life history of Diphyllobothrium erinacei.

(iv) Both poikilothernlic and homoiothermic hosts, viz. tadpoles
and adults of Hyla latopalmata (Gunther) and laboratory-bred mice and rats,
were shown experimentally to serve as reservoir hosts for spargana of
Diphyllobothrium erinacei.

(v) It is postulated that the spargana recorded from Queensland
hosts, as in the list already given, are all of Diphyllobothrium erinacei.

(vi) The adult of Diphyllobothrium erinacei has been shown experi-
mentally to live almost nineteen months within the intestine of a cat (Felis
domestica).

ADDENDUM.

In April, 1952, spargana were recovered from the muscles under the
right arm of a Tasmanian Tiger Cat, Dasyurops maculatus. This host,
from near Launceston, Tasmania, had been flown to Brisbane on dry ice.
The spargana were fed to a cat, and the adult tapeworm, Diphyllobothrium
erinacei , was recovered.

The body of the Tiger Cat was made available for examination through
the kindness of Mr. G. Mack, Director of the Queensland Museum.

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

REFERENCES.

Bearup, A. J. (1948). Observations on the Life Cycle of Diphyllobothrium (Spirometra)
erinacei in Australia (Cestoda : Diphyllobothriidae) . Aust. J. Sci., 10, 183.

Cobbold, T. S. (1879). Parasites ; a treatise of entozoa of man and animals, including
some account of the ectozoa. London xi + pp. 508.

Galliard, H., et Ngu D-V. (1946). Particularity du Cycle I^volutif de Diphyllo-
bothrium mansoni au Tonkin. Ann. Parasit. Hum. Comp., 21, 246.

Gordon, H. McL. (1939). The occurrence of Diphyllobothrium latum, the Broad Fish
Tapeworm, in Dogs in Australia. Aust. Vet. J., 15, 256.

Gan, K. H. (1949). Research on the Life History of Diphyllobothrium ranarum. Docum.
Neerl. Indones. Morb. Trop., 1, 90.

Maplestone, P. A., and Southwell, T. (1923). Notes on Australian Cestodes. Ann.
Trop. Med. Parasit., 17, 317.

Neveu-Lemaire, M. (1936). Traite d’Helminth Medicale et Veterinaire. Nogie
Paris, Vigot, pp. 1514.

Pullar, E. M., (1946.) A Survey of Victorian Canine and Vulpine Parasites. III.

Platyhelminthes other than Taenia multiceps, Taenia ovis and Echinococcus
granulosus. Aust. Vet. J., 22, 40.

Pullar, E. M. (1946). The Control of Internal Parasites in Dogs. Aust. Vet. J., 22, 204.

Pullar, E. M., and McLennan, G. C. (1949). Sparganosis in a Victorian Pig. Aust.
Vet. J., 25, 302.

Sandars, D. F. (1951). Diphyllobothrium latum (Linne) in Australia. Med. J. Aust.,
2, 533.

Wardle, R. A., McLeod, J. A., & Stewart, I. E. (1947). Liihe’s Diphyllobothrium
(Cestoda). J. Parasit., 33, 319.

Young, M. R. (1939). Helminth Parasites of Australia. Imp. Bur. Agric. Parasitol.
(Helminthology). England, pp. 145.

EXPLANATION OF PLATES.

Plate II.

Spargana of Diphyllobothrium erinacei (Rudolphi) in situ in the host Natrix mairii

(Water Snake).

Fig. I. Spargana between the skin and muscles of the body wall, X5*6. Fig. 2.
Spargana in the body cavity between the body walls and gut, X5-6. Fig. 3. Body of
a Water Snake pinned out to show a very heavy infection of spargana, X2-8.

Plate III.

Stages in the Life History of Diphyllobothrium erinacei (Rudolphi).

Fig. 1. Eggs in faeces of an infected cat, X90. Fig. 2. Cyclops varicans
Sars with procercoid larval forms in the thorax and abdomen, X90. Figs. 3 and 4.
Tadpoles of Hyla latopalmata (Gunther) with plerocercoid larval forms (spargana)
distorting both the body and tail regions of the host, X5.

Proo. Roy, Sog, Q’land, Vol. LXIIL, No. 5.

Plate III

Vol. LXIIL, No. 6.

TWO NEW METASTRONGYLE LUNG-WORMS
FROM AUSTRALIAN MARSUPIALS.

By M. Josephine Mackerras and Dorothea F. Sandars*, Queensland
Institute of Medical Research, Brisbane.

(With 6 Text-figures and Plates IV — -VI.)

(. Received 20 th December, 1951; issued separately 6th July, 1953.)

During the last two years forty-five specimens of the common, short-
nosed bandicoot, Isoodon obesulus Shaw and Nodder, have been examined
for parasites. The bandicoots were collected in various Brisbane suburbs
and at Mount Nebo, Mount Tamborine, Nambour, and Gympie. Lung-
worms were found in three specimens, all of which came from Indooroopilly,
a suburb of Brisbane.

Two marsupial mice, Antechinus flavipes Waterhouse, were examined
during the same period. One, which was collected at Mount Glorious, had
a heavy infestation of lung- worms.

As far as can be determined, there are no previous records of lung-
worms in marsupials in Australia, although Travassos (1925) and (1946)
described two lung- worms from South American opossums. The parasites
recovered from the bandicoots and the marsupial mouse are sufficiently
different from each other and from previously described species to warrant
the erection of two new genera, for which the names Marsupostrongylus
and Plectostrongylus are proposed.

The worms were fixed in hot 70% alcohol and preserved in
70% alcohol with 5% glycerine. Unless otherwise stated, examinations
and measurements were made in lacto-phenol.

MARSUPOSTRONGYLUS n. gen.

generic diagnosis : Metastrongylidae with delicate cuticle, which

may be thrown into minute irregular ridges ; no buccal cavity, oesophagus
very short, clavate ; intestine wide. Male with very small bursa, rays
much reduced in size, anterior r&y bilobed, lateral trilobed, externo-dorsal
single, dorso-dorsal ray absent or represented by papillae. Spicules equal
and similar, ending distally in membranous expansions ; gubernaculum
absent. Female with vulva immediately in front of anus, vagina short,
with moderately developed muscular wall. Ovo viviparous. Parasites
of the lungs of marsupials.

type species : Marsupostrongylus bronchialus n. sp.

Marsupostrongylus is perhaps most nearly related to Heterostrongylus
Travassos, from the lungs of South American opossums. It differs from it
in having the spicules equal, in lacking a gubernaculum and in the absence
of a dorso-dorsal ray. The bursa of Heterostrongylus is quite large, with
well- developed dorsal rays, whereas in Marsupostrongylus it is small, with
externo-dorsal rays short and the dorso-dorsal ray rudimentary.

It differs from Plectostrongylus n. gen. in general body form, in the
strongly ornamented cuticle, in the blunt posterior end of the female, in
the short vagina without a strongly developed ovijector, and in the absence
of a gubernaculum in the male.

* And Department of Social and Tropical Medicine, University of Queensland.

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

MARSUPOSTRONGYLUS BRONCHIALUS n. sp.

host : Isoodon obesulus Shaw and Nodder, from Indooroopilly near
Brisbane, South Queensland.

Holotype male and allotype female in the collections of the Queensland
Museum.

habitat : The worms were orientated in the same way in all three

infected bandicoots. They lay parallel to each other in the primary
bronchus of each lung, with their posterior ends free in the lumen and
their anterior ends penetrating the lung. As the primary bronchus was
opened up, it was seen that some worms had entered each secondary
bronchus, and as these in turn were opened up it was found that finally
each female had penetrated singly into a bronchiole. The sharply tapered
anterior end of each worm was pushed into a very fine bronchiole so that
its head came to lie near the pleural surface of the lung.

The intestine of each worm was filled with brownish-black altered
blood. Seen through the transparent cuticle, these dark tubes contrasted
strongly with the milky- white uterine tubes (Plate IV). The males were
usually lying free in the bronchi alongside the females, and their intestines
were also filled with altered blood.

male: Length, 9 to 12 mm.; maximum width, 0*32 mm. at about

2 or 3 mm. from the anterior end. The body tapers towards each end,
the width at the mouth is 0*036 mm., at the oesophageal-intestinal junction,
0*1 mm. and at the cloaca, 0*06 mm. The posterior end is slightly curved
ventrally. The cortical layer of the cuticle is extremely delicate and
voluminous. It has a tessellated appearance due to a mosaic of minute,
irregular ridges. Some of these ridges have a definite transverse trend,
giving the appearance of cross -striations in optical section. This layer is
very loosely attached and is readily distorted during fixation and clearing.
Labial and cephalic structures are inconspicuous ; there appear to be 3
minute lips and 6 small papillae. The mouth leads into the oesophagus,
which is slightly club-shaped and measures 0*23 mm. in length by 0*05 mm.
in maximum breadth (cf. $ Plate V, figs. 1 and 2). The testis is coiled
near the anterior end, and the male duct passes back parallel to the
intestine.

The bursa is small and delicate, with very short, stumpy rays (Plate
V, fig. 5) ; the ventro- ventral and latero- ventral arise from a common
trunk ; the base of the lateral lobe is broad and relatively thick ; the
antero-lateral and postero-lateral rays appear as papilliform projections
on either side of the medio-lateral ray ; the externo- dorsal is single and
rather slender ; the dorso-dorsal ray is apparently represented by two
papillae, one on either side near the base of the externo-dorsal ray ; between
these papillae there appears to be a row of four ill-defined papillae (Text-
fig. 2).

The spicules are similar in size and shape measuring about 0*11 mm.
by 0*01 mm. in maximum breadth. They are brown, narrow and curved.
Proximal] y each ends in an irregular knob, while distally it expands into
a membranous tip supported by three very delicate struts (Plate V, figs.

3 and 6). Neither gubernaculum nor telamon was detected.

Female: Length, 20 to 35 mm.; maximum width, 0*8 to 1*0 mm.
The body is fusiform, tapering very markedly anteriorly so that the anterior
end somewhat resembles a well-sharpened lead pencil. The diameter
at the mouth is 0*05 mm., at the oesophageal -intestinal junction, 0*19

TWO NEW METASTRONGYLE LUNG-WORMS.

mm. The width then increases very rapidly to 0-8 or 1*0 mm. at the region
of the first uterine coils, which lie about 4 mm. from the anterior end.
There is a distinct bulge at this point. The body then gradually narrows
posteriorly, being about 0*5 mm. wide for the greater part of its length.
It decreases to 0*27 mm. at the level of the junction of the uteri, and to
0*11 mm. at the vulva. The vulva is situated immediately in front of the
anus which is 0-06 mm. from the rounded posterior end (Plate V, fig. 4 ;
text-fig. 1).

The cuticle, mouth and oesophagus are similar to those of the male ;
the oesophagus measures 0*23 to 0*27 mm. by 0-05 to 0*06 mm. in maximum
section. The nerve ring appears to be a little anterior to the mid-point
of the oesophagus. The ovaries begin anteriorly and are thrown into several
coils. The uteri, which are packed with developing ova, pass posteriorly
parallel to the intestine. The uteri unite at about 0*3 mm. from the vulva.
The vagina is relatively short, with moderately muscular walls. Ovovivi-
parous.

first stage larva in uterus of female: Length, 0*23 mm. by 0*015
mm. in maximum diameter. There is a slender buccal cavity which leads
into the oesophagus. This is 0*105 to 0*115 mm. long ; it widens slightly
posteriorly where it is about 0*006 mm. in diameter. The intestine and
anus are well differentiated. The nerve ring lies 0*06 mm. from the anterior
end, the tail ends in a sharply pointed, spur-like knob (Text-fig. 3).

Text-figs. 1-3. Marsupostrongylus bronchialus n. sp. 1. Posterior end of female
2. Male bursa ; 3. First-stage larva.

Text-figs. 4-6. Plectostrongylus fragilis n. sp. 4. Posterior end of female , 5. Male
bursa ; 6. First-stage larva, a. anus ; a.l. antero-lateral ray ; d. dorsal papilla ; e.d.
externo-dorsal ray ; i. intestine ; l.v. latero -ventral ray ; m.l. medio-lateral ray ;
o. oesophagus ; p.l. postero-lateral ray ; u. uterus ; v. vagina ; vu. vulva ; v.v.
ventro -ventral ray.

74 PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

PLECTOSTRONGYLUS n. gen.

generic diagnosis: Metastrongylidae with smooth cuticle ; body

filiform ; no buccal cavity ; oesophagus short, simple, and slightly wider
posteriorly. Male with very small bursa, rays much reduced in size, ventral
ray bilobed, lateral trilobed, externo- dorsal single, dorso-dorsal ray absent
or represented by papillae. Spicules equal and similar, ending distally
in membranous expansions. Gubernaculum present. Female with posterior
extremity straight, tail pointed ; ovijector well developed ; vulva anterior
to anus and close to it. Ovo viviparous. Parasites of the lungs of marsupials.

type species : Plectostrongylus fragilis n. sp.

Plectostrongylus resembles Marsupostrongylus in the bursal formula
and having the spicules equal. It differs from it in the possession of a
gubernaculum in the male ; in the female the tail is pointed, and there is a
long vagina ending in a muscular ovijector, whereas in Marsupostrongylus
the tail is blunt and the vagina is relatively very short, with moderately
muscular walls.

It differs from Heterostrongylus in having a greatly reduced bursa,
in the absence of the dorso-dorsal ray, and in having the spicules equal
and similar.

In the extreme reduction of the dorsal ray it seems to resemble
Pneumostrongylus Monnig from the impala, but differs from it in the reduced
bursa, the bursal formula, the presence of a gubernaculum, the absence
of a telamon. The eggs of Pneumostrongylus are segmenting when laid,
whereas Plectostrongylus is ovo viviparous.

PLECTOSTRONGYLUS FRAGILIS n. sp.

host : Antechinus flavipes Waterhouse, from Mount Glorious, South

Queensland.

Holotype male and fragments of females from the same host in the
collection of the Queensland Museum.

habitat: The worms lay in the bronchioles, including the finer ones
(Plate VI, fig. 7), and some even invaded the alveoli (Plate VI, fig. 6). The
generic name indicates the intimate manner in which the worm is woven
into the tissue of the lung.

male. One intact specimen, one complete but broken specimen and
several fragments were obtained. The body is filiform, length 10 to 15
mm. by 0-088 mm. to 0*114 mm. in maximum diameter. The width at
the oesophageal-intestinal junction is about 0*042 mm. Labia and cephalic
papillae are inconspicuous. The cuticle is smooth. The mouth leads into
the oesophagus, which is short, simple and slightly club-shaped, measuring
0*244 mm. in length by 0*02 mm. in maximum width. The intestine is a
narrow tube.

The bursa is very small, and the rays are short and stumpy with a
tendency to appear pedunculated (Plate VI, fig. 4). The ventro- ventral
and latero-ventral arise from a common trunk, the ventro-ventral being
the larger ; these rays are incurved in all our specimens, but are shown
diagrammatically as pointing outward in text-fig. 5. The antero-lateral
and postero-lateral are broad and slightly larger than the medio-lateral ;
all the laterals arise from a common trunk ; the externo -dorsal is short
and blunt ; the dorso-dorsal ray appears to be represented by two small
papillae, one on each side near the base of the externo -dorsal ray.

TWO NEW METASTRONGYLE LUNG-WORMS.

The spicules are equal and similar. The proximal portion of each is
broad, and the distal portion is drawn out into two fine rods, each sur-
rounded by a membranous expansion. The spicules measure 0*09 to 0*1
mm. in length by 0*01 mm. in maximum breadth ; the membranous part
is about 0*04 mm. long (Plate IV, figs. 2, 3 and 5). A V -shaped gubernaculum
is present (Plate IV, fig. 2).

female. No intact specimens were recovered, so the length could
not be determined. However, the fragments which were obtained suggested
that the females are considerably longer than the males. The maximum
width of some of the fragments is 0*1 mm. The width at the vulva is 0-04
mm. and the anus 0-026 mm. The posterior end is obliquely truncated,
and the tail sharply pointed. The anus lies 0-036 to 0-05 mm., and the
vulva 0*09 to 0-1 mm. from the tip of the tail (Text-fig. 4).

In one specimen, fixed in formol-acetic-alcohol and stained with
haematoxylin, the vagina measures 0-83 mm. in length and terminates
in a muscular ovijector. The uteri and vagina are packed with ova con-
taining well-developed embryos. The ova measure 0-05 to 0-055 mm.
by 0-026 to 0-03 mm.

first-stage larva in uterus of female: Length 0-19 to 0-20 mm.
by 0-008 to 0-01 mm. in width. There is a slender buccal cavity leading
into the oesophagus, which is 0-09 to 0-095 mm. in length. The nerve ring
is situated 0*05 mm. from the anterior end. The tail is sharply pointed
(Text-fig. 6 ; Plate VI, fig. 1).

DISCUSSION.

The metastrongylid lung-worms have been studied recently by
Dougherty, who has proposed a classification and discussed the evolution
of the group (Dougherty, 1949, 1951). He has divided the family into six
sub-families : — Metastrongylinae, Filaroidinae, Skrjabingylinae, Pseuda-
liinae, Protostrongylinae and Dictyocaulinae. The Filaroidinae (with
six genera) have developed mainly in the Carnivora, the Pseudaliinae
(four genera) in the Cetacea and the Protostrongylinae (ten genera) in the
Artiodactyla. The Dictyocaulinae contains only one genus, Dictyocaulus,
parasitic in the Ungulata, and bearing some striking resemblances to the
Trichostrongylidae. The Skrjabingylinae (5 genera) occur in the Carnivora ,
Insectivora and one species, Troglostrongylus delicatus Travassos 1946,
in the South American opossum . Dougherty places Heterostrongylus , the only
other genus known from marsupials, in the sub -family Metastrongylinae,
together with Metastrongylus which is parasitic in pigs. Heterostrongylus
heterostrongylus Travassos 1925 was described from the lungs of the South
American opossum Didelphis marsupialis aurita Wied. It possesses a
fairly large bursa with well- developed rays, although the arrangement of
the dorsal system is unusual.

The two new genera described here cannot be definitely allotted to
any of the above sub -families, although they appear to resemble some of
the filar oidine genera in the reduction of the bursa. We think that, until
more material is studied from Australian marsupials and some of the life-
histories are elucidated, it is wisest to refrain from putting these genera
into any of the known sub -families.

The hosts of our new species are polyprotodont marsupials belonging
to different families. Isoodon belongs to the Peramelidae, which includes
the bandicoots and rabbit-bandicoots, ground- dwelling, insectivorous or
omnivorous creatures. Antechinus belongs to the Dasyuridae, which includes

PROCEEDINGS OF THE ROYAL SOCIETY OF QUEENSLAND.

the carnivorous marsupials, the smaller members being mainly insecti-
vorous. The South American opossums are also polyprotodonts, belonging
to the family Didelphidae, arboreal creatures, mainly insectivorous or
carnivorous.

It may be that the relationships of the parasites in the Metastrongylidae
are a reflection of the habits of the hosts rather than of their phylogeny.

ACKNOWLEDGEMENTS.

We are indebted to Dr. E. Singer of this Institute for bringing us
numerous bandicoots, including two of the infected specimens, and to
Mr. G. Naylor for the other infected specimen. We also would like to
thank Mr. G. Mack of the Queensland Museum for identifying the marsupial
hosts.

SUMMARY.

Two new metastrongylid lung- worms are described, Marsupo-
strongylus bronchialus n. gen., n. sp., from the bandicoot, Isoodon obesulus ,
and Plectostrongylus fragilis n. gen., n. sp., from a marsupial mouse,
Antechinus flavipes.

REFERENCES.

Dougherty, E. C. (1949). The Phylogeny of the Nematode Family Metastrongylidae
Leiper, [1909] : a correlation of host and symbiote evolution. Parasitology ,
39, 222-234.

Dougherty, E. C. (1951). A further revision in the classification of the Family
Metastrongylidae Leiper [1909] (Phylum Nematoda). Parasitology, 41, 91-96.

Travassos, L. (1925). Un nouveau type de Metastrongylidae. Comp. rend. Soc. Biol.,
93, 1259-1262.

Travassos, L. (1946). Rev. Brasil. Biol., 6, 499.

EXPLANATION OF PLATES.

Plate IY.

Marsupostrongylus bronchialus n. gen. and sp.

Bronchus of lung of Isoodon obesulus, opened to show the tails of lung-worms
lying free in the lumen with their anterior ends entering the bronchioles. Note white
uterine tubes and dark gut content of the worm.

Plate V.

Marsupostrongylus bronchialus n. gen. and sp.

Fig. 1. Anterior end of body of $. Fig 2. Cephalic region of same $ showing
oesophagus and tessellated cortical layer of the cuticle. Fig. 3. Side view of tail of $.
Fig. 4. Side view of tail of $. Note blood in the gut. Fig. 5. Side view of tail of $
showing the small bursa and rays. Fig. 6. Side view of tail of $ showing the tip of
one spicule, which is extruded.

Scale. In fig. 1, one division of the scale is 0-05 mm. In figs. 2-6 inclusive each
division of the scale is 0 01 mm. Figs. 2 and 4 are to the same scale.

Plate YI.

Plectostrongylus fragilis n. gen. and sp.

Fig. 1. First stage larva in smear made from fragments of an adult female.
Fig. 2. Ventral view of tail of showing the gubernaculum, bursa, extended
membranous tips of spicules. Fig. 3. Lateral view of tail of showing the spicules
separated. Fig. 4. Lateral view of tail of <£, showing bursal rays. Fig. 5. Lateral
view of tail of $, showing spicules. Fig. 6. Section of lung of Antechinus flavipes
showing cross sections of both and 2 adult Plectostrongylus fragilis in the alveoli.
Fig. 7. Section of lung of Antechinus flavipes showing cross sections of both $ and $
adult Plectostrongylus fragilis coiled up in a bronchiole.

The divisions of the scale represent 0 01 mm. in each figure. Figs. 2-5 are to the
same scale ; figs. 6-7 are to the same scale.

Pkoc. Roy. Soc. Q’land., Yol. LXIIL, No. 6.

Plate TV ►

. Roy. Soc. Q’land., Yol. LXIII., No. 6.

Plate V.

CSV *

Proc. Roy. Soc. Q’land.; Vol. LXIII., No. 6.

Plate VI.

The Royal Society of Queensland.

Report of the Council for 1950.

To the Members of the Royal Society of Queensland.

Your Council has pleasure in submitting- the Annual Report of the
Society for the year 1950.

At Ordinary Meetings throughout the year five addresses were
given, while one evening was devoted to films, another to a short film
followed by exhibits, and a third to three short talks.

Several original papers were accepted for publication in the
Proceedings.

The Council has decided to publish a C. T. White Memorial
Supplement to the 1950 Volume of Proceedings. This is at present
being prepared.

The delay in publication has partly been overcome. Volume LXI.
for 1949 is about to be issued, and Volume LXII. is in the press.

Our thanks are due to the Librarian for the excellent work he has
done in the Library. One-third has now been catalogued, and in many
cases, missing numbers in series have been obtained.

The Society has supported the establishment of a Marine Biological
Research Station on the Barrier Reef.

There are now 5 honorary life members, 10 life members, 3
corresponding members, 238 ordinary members and 12 associate members
in the Society. During the year the Society lost 2 members by death
and 7 by resignation;, 17 ordinary members and 13 associate members
have been elected. Mr. L. C. Ball was elected to honorary life
membership.

Attendance at Council Meetings was as follows : — M. F. Hickey, 10 ;
D. Hill, 8 ; H. J. G. Hines, 7 ; M. I. R. Scott, 10 ; D. F. Sanders, 9 ;
F. S. Colliver, 9; S. T. Blake, 9; G. Mack, 9; I. M. Mackerras, 8;
A. L. Reimann, 8 ; J. H. Simmonds, 7 ; L. J. H. Teakle, 7 ; H. C. Webster,
8.

M. F. HICKEY, President.

Margaret I. R. Scott, Hon. Secretary.
21st March, 1951.

VI.

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ABSTRACT OF PROCEEDINGS.

VII.

Abstract of Proceedings, 2nd April, 1951.

The Annual General Meeting of the Society was held in the Physics
, Department of the University on Monday, 2nd April, with the President
(Associate Professor M. F. Hickey) in the chair. About fifty members
and friends were present. The minutes of the last Annual Meeting
were read and confirmed. The Annual Report was adopted and the
Balance-Sheet received. The Librarian reported 584 volumes and parts
had been added to the library and that new exchanges had been
established with the Japanese Academy of Science, the University of
Southern California, and the Rancho Santa Anna Botanical Gardens.

Professor T. K. Ewer and Professor F. T. M. AVhite were elected
to Ordinary Membership.

The following officers were elected for 1951 : —

President : H. J. G. Hines.

Vice-President : I. M. Mackerras.

Hon. Secretary : Miss M. I. R. Scott.

Hon. Treasurer : Miss D. F. Sandars.

Librarian : F. S. Colli ver.

Editors: S. T. Blake, George Mack.

Councillors : M. J. Mackerras, A. L. Reimann, J. H. Simmonds,
W. Stephenson, L. J. H. Teakle.

Hon. Auditor : L. P. Herdsman.

Before delivering his address, the President remarked as follows : —

Since the last Annual General Meeting, we have suffered the great
loss of two distinguished members, Mr. C. T. White and Mr. J. B.
Henderson. It is with deep regret that I recall them to your memories
to-night.

Mr. White, Government Botanist at the time of his death, died in
harness. For a long time a member of our Society and of the Council
of the Society, he was as well-loved personally as he was distinguished
scientifically. His reputation was international.

Mr. Henderson, formerly Chief Government Analyst and an
outstanding member of the Public Service of Queensland, was a very
old member of our Society and one held in the highest esteem. He was
a trustee of the Society.

Before turning to the main part of my address I wish to express
my thanks to the members of the Council for the pleasant term of
office I have enjoyed. I think that the Council would wish me to
express appreciation of the work done by the Honorary Librarian,
Mr. Colliver, in the reorganization of the Library, the most precious
material asset of the Royal Society of Queensland.

The Presidential Address, entitled “Form or Function,” was
delivered by Associate Professor M. F. Hickey.

Abstract of Proceedings, 7th May, 1951.

The Ordinary Monthly Meeting of the Society was held in the
Physics Department of the University on Monday, 7th May, with the
President (Associate Professor H. J. G. Hines) in the chair. About
sixty members and friends were present. The minutes of the previous

VIII.

ABSTRACT OF PROCEEDINGS.

meeting were confirmed. The following were nominated for
membership : — Mr. A. A. Gibson, Mr. R. N. Spratt, Mr. A. W. Draydon,
Dr. F. G. Christensen and Dr. W. Hayhow for Ordinary Membership,
and Miss M. Lewis for Associate Membership. Mr. F. Gipps was
elected to Honorary Life Membership.

Dr. I. M. Mackerras gave a brief account of an outbreak of
encephalitis in the Murray Valley, Victoria. This had been found by
workers at the Hall Institute to be closely related to the Japanese B
type of encephalitis. A much milder outbreak had occurred among
natives at Mornington Island during February and March. Preliminary
serological studies at the Hall Institute had shown that this was closely
related to the Murray Valley infection. A full serological investigation
was being made at Mornington Island in association with a search for
the insect vectors and animal reservoirs of the infection.

Professor H. J. Wilkinson gave an address entitled “Prehistoric
Settlement of the Pacific Islands,” He said that the Pacific Islanders
are of at least three distinct types, the Melanesians, Polynesians and
Micronesians, and each type is the result of the mixture of two or more
of the basic racial groups. Melanesians are mostly negroid, but
several other racial elements are present and vary in intensity in
different parts of the Melanesian islands, Polynesians are patently a
hybrid race compounded of white, mongoloid and a modicum of negroid.
The Micronesians are rapidly disappearing and are not very well
known, but they seem to have been allied to the Polynesians in race
and culture, though with strong Indonesian mongoloid influences.
It was shown how Physical Anthropology, blood grouping investigations,
Ethnology, the study of languages, customs and traditions, etc., as well
as of the distribution of plants and animals, are all contributing
towards the solution of the problem of the origin of the Pacific
Islanders and the diffusion of culture in the Pacific. Comments were
made on the recent book by Gladwyn “Men out of Asia” and on the
significance of Hayerdahl’s “Kon Tiki Expedition.” The lecture was
profusely illustrated with lantern slides.

Abstract of Proceedings, 25th June, 1951.

The Ordinary Monthly Meeting of the Society was held in the
Physics Department of the University on Monday, 25th June, with
the President (Associate Professor H. J. G. Hines) in the chair. About
thirty members and friends were present. The minutes of the previous
meeting were confirmed. The following were elected to Membership : —
Mr. A. A. Gibson, Mr. R. N. Spratt, Mr. A. W. Draydon, Dr F. G.
Christensen and Dr. W. Hayhow to Ordinary Membership ; and
Miss M. Lewis to Associate Membership.

Dr. Owen Jones exhibited a part of the lower and upper jaws with
well preserved teeth of Ichthyosaurus australis McCoy, from near
Richmond, Central Queensland. The specimen was sent to Professor
S. F. Lumb who passed it on to the Geology Department of the
University.

ABSTRACT OF PROCEEDINGS.

IX.

Professor W. H. Bryan exhibited a new geological map of
Brisbane on a scale of 1 mile to an inch, prepared by himself and
Dr. Owen Jones. It shows much more detail of rock type and
structure than earlier maps.

Professor W. H. Bryan exhibited a fine specimen from the quarry
in the Brisbane Tuffs at Stafford. The specimen is of a rhyolitic
agglomerate with large fragments of phyllite. It was probably derived
from the throat of a nearby volcano.

Research Professor A. L. Reimann exhibited a new type of
monochromatic optical filter, a Barr and Stroud “interference filter,”
having a narrow transmission band with its maximum transmission at
the wave-length of the green mercury line, viz., 5461 A°. The location
and width of the transmission band were shown by projecting a
spectrum on a screen and interposing the filter in the path of the
light. An outline of the theory and method of construction of this
filter was given, it being pointed out that it is in principle nothing
more than a logical development of the Fabry and Perot etalon, which
was devised as long ago as 1901.

Mr. W. B. Mather exhibited four strains of Drosophila
melanog aster, viz., wild type, white eye, ebony body, and ebony body-
dumpy wing, used in the senior genetics course of the Zoology
Department. He pointed out that Drosophila is ideal for cytogenetical
work because it is easy to breed, has a short life cycle, has a low
chromosome number, and forms giant salivary gland chromosomes.
The white eye mutant is used for illustrating sex linkage and criss-
cross inheritance, the ebony body for the monohybrid ratio, and the
ebony body-dumpy wing for the dihybrid ratio.

Mr. George Mack exhibited two specimens, a marsupial, the brown
cuscus ( Phalanger orient alis) , and a eutherian mammal, the giant rat
( Uromys caudimaculatus) . Both species are confined to Cape York
Peninsula. The brown cuscus was discovered in Australia only in
recent years, and the need for improved knowledge of the fauna was
stressed in view of the fundamental importance of the indigenous flora
and fauna in relation to soil erosion. Mr. Mack referred to the
tendency to treat soil erosion as a mechanical problem, whereas, he
insisted, it is primarily a biological problem.

Dr. I. M. Mackerras, Dr. M. J. Mackerras and Miss D. F. Sandars
exhibited parasites of the bandicoot ( Isoodon obesidus).

Abstract of Proceedings, 30th July, 1951.

The Ordinary Monthly Meeting of the Society was held in the
Physics Department of the University on Monday, 30th July, with the
President (Associate Professor H. J. G. Hines) in the chair. About
thirty members and friends were present. The minutes of the previous
meeting were confirmed. Miss B. J. Excell was nominated for Ordinary
Membership. 395 volumes and parts have been added to the library.

A symposium was held on “The Zoology of Copper.” Mr. J. M.
Harvey and Mr. J. E. 0 ’Hagan were the principal speakers.

ABSTRACT OF PROCEEDINGS.

Mr. Harvey gave a brief historical account of copper deficiency
in ruminants, and then showed how copper administration had been
attempted. There were five methods commonly used — (i.) Through
the pastures by top dressing of the soil with copper salts, (ii.) Through
the drinking water, (iii.) By oral administration as drenches, (iv.)
By addition of copper compounds to exposed mineral supplements,
(v.) By direct incorporation in the food.

All suffered certain disadvantages arising from either cost, lack
of controlled intake, inactivation, or simply from impracticability.
In the work he outlined, attempts were made to establish within the
animal ample reserves of copper from which the necessary small
continual releases could take place. The methods used were —
(a) Oral administration, (b) Intramuscular injection, (c) Subcutaneous
injection, (d) Implantation, and the preparations used were organic
and inorganic compounds of copper of both soluble and insoluble types.
All were either readily available, or presented no great difficulty in
preparation. Balance sheets were kept to assess copper retention.
Though some untoward experiences — abscesses, necrotic areas and even
deaths — were recorded, it was eventually possible to employ intra-
muscular injection of soluble copper salts of simple inorganic and
organic acids, and implantation of the copper compound of Hahn’s
oxine reagent (8 hydroxyquinoline). These are promising and large
scale field trials are envisaged.

Mr. 0 ’Hagan said that with one exception, the copper uroporphyrin
complex turacin first found in the feathers of certain African birds,
copper occurs in plant and animal tissues in combination with proteins.
These copper proteins can be divided into three classes (i.) the plant
and animal oxidases, (ii.) the haemocyanins, (iii.) the animal copper
proteins of blood, liver, etc. The oxidases, laccase, tyrosinase and
ascorbic acid oxidase, are enzymes responsible for the respiratory
activity of many plants. Evidence of their activity is the darkening of
many cut plants (such as the sweet potato tuber and the banana) on
contact with air. The haemocyanins, the respiratory pigment of the
blood of crabs, lobsters, octopi, snails, certain worms and molluscs have
molecular weights varying from 350,000 to 5,000,000, contain two atoms
of copper per molecule and can carry up to 25 ml. of oxygen per 100 g.
of protein. They resemble the haemoglobins but have copper in place
of iron. The animal copper proteins, haemocuprein and hepatocuprein
occur respectively in the blood and liver of higher animals and man.
These proteins are responsible for the storage, transport and perhaps
the activity of copper as a necessary component of the system or
systems responsible for the synthesis of haemoglobin, cytochrome and
cytochrome oxidase. It was shown that copper plays an essential role
in the respiratory activity of plants, animals and men.

Abstract of Proceedings, 27th August, 1951.

The Ordinary Monthly Meeting of the Society was held in the
Physics Department of the University on Monday, 27th August, with
the President (Associate Professor H. J. G. Hines) in the chair. About
forty-five members and friends were present. The minutes of the
previous meeting were confirmed. Miss B. J. Excell was elected to

ABSTRACT OF PROCEEDINGS.

XI.

Ordinary Membership. Mr. R. Millar was nominated for Ordinary
Membership. The Librarian reported that sixty-six volumes and parts
had been added to the Library during August, and that a new exchange
had been established with the Hungarian Academy of Science.

Professor Schonell gave an address entitled “Psychology in
Educational Practice.” A lengthy discussion followed.

Abstract of Proceedings, 24th September, 1951.

The Ordinary Monthly Meeting of the Society was held in the
Physics Department of the University on Monday, 24th September,
with the President (Associate Professor H. J. G. Hines) in the chair.
About thirty-five members and friends were present. The minutes of
the previous meeting were confirmed. Mr. R. Millar was elected to
Ordinary Membership. The Librarian reported that 130 volumes and
parts had been added to the Library since the last meeting, and that
a new exchange had been established with the Finnish Academy of
Science.

Mr. James J. Bollich spoke on “The Relationship of Gravity,
Magnetism and Atmospheric Electricity to Tectonics,” He said that
as more and more data concerning the phenomena of gravity, magnetism,
atmospheric electricity, earth-currents, and earthquakes are accumulated
it is becoming increasingly apparent that these phenomena are very
closely related. If this relationship exists it then seems reasonable to
assume that these phenomena must all originate from a single cause.
The speaker proposed a hypothesis based upon differential rotation
between crustal and subcrustal material of the earth, comparable to the
difference in the speed of rotation of the two mediums of a revolving
glass of water, to explain these phenomena. This mechanism has been
advanced previously to explain magnetism and atmospheric electricity
and the speaker believes that it should be investigated further as the
force responsible for crustal deformation.

Dr. R. Gradwell spoke on “Petrological Research at Imperial
College, London.” He described research being carried on at the
Geology Department of the Imperial College of Science and Technology
by Professor H. H. Read and his staff and students. Most of this
research concerns Plutonic rocks, and it was pointed out that great
importance is attached to field geology. Ironstones from Wellingborough,
England, were exhibited and some colour slides of the open-cut workings
were shown. Shap and Cornish granites were also exhibited.

Dr. Owen Jones informed members that at a meeting held in May
and representative of the Geologists of Australia it was decided to
form a Geological Society of Australia. He gave an outline of the
proposed constitution and explained that the main object of the
Society was to publish papers of a high standard, particularly those
of world- wide interest and ones which are too long for the journals
of existing societies.

Dr. Dorothy Hill exhibited R. van Bemmelen’s “Geology of
Indonesia,” a work in two volumes with a case of maps, which sets
out the results of 100 years’ work, mainly by the Dutch, in Indonesia,
and interprets the findings in the light of his theories on the relation
between earth movement and igneous activity.

XII.

ABSTRACT OF PROCEEDINGS.

Associate Professor F. W. Whitehouse exhibited new early
Carboniferous fossils from Newry Island, and a new genus of early
Cretaceous lamellibranchs from near Tambo.

Mr. Grahame Tweedale exhibited some specimens of slate green-
stone, phyllite and altered basic lavas from the Gogango Range, between
Rockhampton and Duaringa, introducing the exhibit with a few remarks
on the geological structures of the area. The Gogango High divides
the Yarrol Basin, with its Devonian, Carboniferous and Permian
sediments, from the Permo-Carboniferous Bowen Basin to the west.
Slates of the type exhibited are very rare in Queensland, and a further
interesting feature is their association with regionally metamorphosed
schists, phyllites and greenstones.

Abstract of Proceedings, 29th October, 1951.

The Ordinary Monthly Meeting of the Society was held in the
Physics Department of the University on Monday, 29th October, with
the President (Associate Professor H. J. G. Hines) in the chair.
About twenty members and friends were present. The minutes of the
previous meeting were confirmed. Mr. J. E. Kindler was nominated
for Ordinary Membership. The Librarian reported 184 additions to
the library, and the establishment of a new exchange for Memoirs of
Das’ A’lvaro de Castro Museum, since the last meeting.

Dr. M. C. Bleakly gave an address entitled “Some Features of the
Development of Callorhynchus milii (Elephant Fish) having a bearing
on the Primitiveness of the Holocephali. ” Three points from the
development of Callorhynchus milii were considered. A study of the
development of the jaw suspension, of the relations of the spiracle up
to the time of its closure, and of the development of the hyoid skeleton
supports the view that the complete hyoid skeleton, unique amongst
living vertebrates, is a retained primitive feature and not a secondary
modification. The kidney shows features more primitive than that of
any jawed vertebrate since all the segments from which it is developed
are retained, are recognisable and are functional in the adult. . The
relations of the mandibular arterial arches to the trabeculae of the
developing skull are as in all vertebrates save the Selachii which
therefore appear to be a modified group. These features support the
view put forward by de Beer that the Holocephali are not derived from
the Selachii but represent the more primitive living Gnathostomes
known.

Abstract of Proceedings, 26th November, 1951.

The Ordinary Monthly Meeting of the Society was held in the
Physics Department of the University on Monday, 26th November,
with the President (Associate Professor PI. J. G. Hines) in the chair.
About fifty-five members and friends were present. The minutes of
the previous meeting were confirmed. Mr. J. E. Kindler was elected
to Ordinary Membership. The Librarian reported the addition to the
library of il9 volumes and parts for November.

ABSTRACT OF PROCEEDINGS.

XIII.

A Symposium on “Aerial Photographic Interpretation” was held.

Mr. Newell explained the general principles behind the taking of
aerial photographs, the principles of stereoscopy and various aspects,
optical and cartographic, in the preparation of maps for aerial
photographs.

Mr. Reilly, with the aid of lantern slides, described the equipment
used to-day in the Department of the Surveyor-General for the making
of topographic maps from aerial photographs.

Major Herridge (D.A.D. Survey), exhibited the several types of
maps prepared by the Army for aerial photographs and discussed the
techniques employed.

Mr. Woods illustrated the use to which aerial photographs are put
by the Queensland Forest Service for making special maps for
delimiting the distribution of timbers of economic importance.

Mr. Tweedale discussed pattern mapping and the uses that are
being made of it at present in preparing a new Geological Map of
Queensland.

Associate Professor Whitehouse, with the aid of lantern slides of
aerial photographs, demonstrated a variety of special applications —
road locations, soil survey, coastal problems, etc.

A. H. Tucker, Government Printer, Brisbane.

GUIDE FOR THE PREPARATION OF SYNOPSES

1. PURPOSE.

It is desirable that each paper be accompanied by a synopsis preferably
appearing at the beginning. This synopsis is not part of the paper; it is intended
to convey briefly the content of the paper, to draw attention to all new information
and to the main conclusions. It should be factual.

2. STYLE OF WRITING.

It should be presumed that the reader has some knowledge of the subject
but) has not read the paper. The synopsis should therefore be intelligible in itself
without reference to the paper, for example it should not cite sections or illustra-
tions by their numerical references in the text.

3. CONTENT.

It is sometimes valuable to indicate the treatment of the subject by such
words as: brief, exhaustive, theoretical, etc.

The synopsis should indicate newly observed facts, conclusions of an experiment
or argument and, if possible, the essential parts of any new theory, treatment,
apparatus, technique, etc.

It should contain the names of any new compound, mineral, species, etc., and
;any new numerical data, such as physical constants; if this is not possible it should
draw attention to them. It is important to refer to new items and observations,
*even though some are incidental to the main purpose of the paper; such) information
may otherwise be hidden though it is often very useful.

When giving experimental results the synopsis should indicate the methods
used; for new methods the basic principle, range of operation and degree of
accuracy should be given.

4. DETAIL OF LAYOUT.

It is impossible to recommend a standard length for a synopsis. It should,
however, be concise and should not normally exceed 100 words.

If it is necessary to refer to earlier work in the summary, the reference should
always be given in the same manner as in the text. Otherwise references should
be left out.

CONTENTS

vol. lxiil

No. i, — Form or Function. By M. F. Hickey. (Issued separately,

29th June, 1953)

No. 2. — Studies of the Life Histories of Some Queensland Blattidae

(Orthoptera). Part 1. The Domestic Species. By Pauline

Pope. (Issued separately, 6th July, 1953)

No. 3.— Studies of the Life Histories of Some Queensland Blattidae

(Orthoptera). Part 2. Some Native Species. By Pauline

Pope. (Issued separately, 6th July, 1953) .. • • ••

No. 4. — Parasites of the Bandicoot, Isoodon oltesulus. By I. M. Mackerras,
M. J. Mackerras and D. F. Sandars. (Issued separately,

6th July, 1953) •

No. 5. — A Study of Diphyllobothriidae (Cestoda) from Australian Hosts.

By Dorothea F. Sandars. (Issued separately, 6th July, 1953)

No.. 6. Two New Metastrongyle Lung-worms from Australian Marsupials.

By M. Josephine Mackerras and Dorothea F. Sandars. (Issued
separately, 6th July, 1953)

Eeport of Council

Abstract of Proceedings

Pages.

1-22

23-46

47-60

61-64

65-70

71-76

vii.

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